Skip to main content

Full text of "Bulletin of the Natural History Museum Zoology"

See other formats


ISSN 0968-0470 



Bulletin of 

The Natural History 

Museum 



Zoology Series 




THE 

NATURAL 
HISTORY 
MUSEUM 



VOLUME 62 NUMBER 1 27 JUNE 1996 



The Bulletin of The Natural History Museum (formerly: Bulletin of the British Museum 
(Natural History)), instituted in 1949, is issued in four scientific series, Botany, 
Entomology, Geology (incorporating Mineralogy) and Zoology. 

The Zoology Series is edited in the Museum's Department of Zoology 
Keeper of Zoology: Dr C.R. Curds 

Editor of Bulletin: Dr N.R. Merrett 

Assistant Editor: Dr B.T. Clarke 



Papers in the Bulletin are primarily the results of research carried out on the unique and ever- 
growing collections of the Museum, both by the scientific staff and by specialists from elsewhere 
who make use of the Museum's resources. Many of the papers are works of reference that will remain 
indispensable for years to come. All papers submitted for publication are subjected to external peer review for 
acceptance. 

A volume contains about 1 60 pages, made up by two numbers, published in the Spring and Autumn. 
Subscriptions may be placed for one or more of the series on an annual basis. Individual numbers and back 
numbers can be purchased and a Bulletin catalogue, by series, is available. Orders and enquiries should be sent 
to: 

Intercept Ltd. 

P.O. Box 716 

Andover 

Hampshire SP 10 1YG 

Telephone: (01264) 334748 
Fax: (01264) 334058 

Claims for non-receipt of issues of the Bulletin will be met free of charge if received by the Publisher within 6 
months for the UK, and 9 months for the rest of the world. 



World List abbreviation: Bull. nat. Hist. Mus. Lond. (Zool.) 



© The Natural History Museum, 1996 



ISSN 0968-0470 

The Natural History Museum 
Cromwell Road 
London SW7 5BD 



Zoology Series 

Vol. 62, No. l,pp. 1-70 



Issued 27 June 1996 



Typeset by Ann Buchan (Typesetters), Middlesex 
Printed in Great Britain at The Alden Press, Oxford 



Bull. nat. Hist. Mus. Lond. (Zool.)62(l): 1-30 



Issued 27 June 1996 



Deep-sea conoidean gastropods collected by the 
John Murray Expedition, 1933-34 

ALEXANDER V. SYSOEV 

Zoological Museum of Moscow State University, Hertzen Street 6, Moscow 103009, Russia 

Synopsis. Conoidean gastropod molluscs (formerly treated within the family Turridae) from 20 deep-sea (bathyal) 
stations of the John Murray Expedition (North-Western Indian Ocean) are described. A total of 50 species from 11 
families and subfamilies, and 3 1 genera and subgenera, have been found in the material studied. 1 7 species are described as 
new. 



INTRODUCTION 



TAXONOMY 



The John Murray Expedition worked aboard the 'Mabahiss' in 
the northwestern part of the Indian Ocean in 1933-34 and 
collected invaluable biological material from the area which has 
not been subsequently explored on such a large scale. However, 
no account of the gastropods collected by that expedition have 
ever been published. Through the courtesy of Dr John D. Taylor 
of The Natural History Museum, London I was able to study the 
conoidean gastropods from that collection. This paper deals 
with deep-sea samples of that part of the Conoidea (= 
Toxoglossa) which was usually treated as the family Turridae. 
The Turridae s.l., unlike the Terebridae and Conus, are very 
characteristic of the deep-sea molluscan fauna, of which they 
comprise a considerable part. The material studied was 
previously loaned to the late A.W.B. Powell who mentioned 
some of the species in two parts of his revision of Indo-Pacific 
Turridae (Powell, 1964, 1969). However, none of Powells species 
identifications was found on labels accompanying the samples. 



MATERIAL AND METHODS 



The molluscs studied were collected at 20 bathyal (depth 
183-2312 m) stations of H.E.M.S. 'Mabahiss' in the 
North-Western Indian Ocean (Table 1). The classification of 
Conoidea used in the present paper follows that adopted by 
Taylor et al. (1993). All the material including the type 
specimens of the new species is stored in the Natural History 
Museum. 



ABBREVIATIONS USED IN THE TEXT 

NHM The Natural History Museum, London 

JME John Murray Expedition 

H shell height 

D shell diameter 

Hs spire height 

Hb body whorl height 

Ha aperture height 



Family DRILLIIDAE Morrison, 1966. ICZN pending 
Genus DRILLIA Gray, 1838 

Type species: Drillia umbilicata Gray, 1838 (subsequent 
designation Gray, 1847) 

Drillia altispira Sysoev, new species 

Figs 18 & 19 

Material, stn 176, 1 shell (holotype, No. 1993088). 

Description. The shell is rather large, claviform, with very 
high spire exceeding half of the shell height, thick and solid, 
light-brown, consisting of almost 12 whorls. The protoconch is 
missing. The whorls are weakly convex and slightly angled; the 
point of angulation is situated below the periphery in the spire 
whorls, but shifts upwards on the last whorls. The subsutural 
slope is concave, and the prominence of concavity increases 
towards the body whorl. Sutures are clear, straight, and shallow. 
Axial sculpture consists of oblique, narrowly crested folds 
terminating on the subsutural slope. Some folds form weak 
nodules just below the suture. The folds become subobsolete on 
the last quarter of body whorl, probably as a result of preceding 
shell damage. There are 14 folds on the penultimate whorl and 
about the same number on the body whorl. Spiral ribs are 
numerous, uniform, rounded, moderately strong, with 
interspaces equal to them in width. The ribs cover the entire shell 
surface but become narrower, closer, and weaker on the 
subsutural slope. The shell base forms a weak bend in passing to 
a moderately developed fasciole. The aperture is rather small, 
inversely pyriform, with a distinct stromboid notch. The outer 
lip with a thin edge, projects strongly and forms an alate 
expansion between the anal sinus and stromboid notch. There is 
no prominent prelabral varix, only a thin fold curved in 
correspondence to growth lines is present. The inner lip is 
covered by thick and wide glossy callus which is mostly free 
along its outer edge and forms a shallow false umbilical cavity. 
The parietal callus pad is large and rounded, constricting the 
entrance to the anal sinus. The anal sinus is deep, U-shaped, with 
spout-like edge, directed slightly adapically. The canal is short, 
slightly bent to the right, shallowly notched and obliquely 
truncated. H = 37.9, Hb = 17.5, Ha = 14.2, D = 1 1 .4 mm. 



©The Natural History Museum, 1996 



A.V. SYSOEV 



Table 1 Stations of H.E.M.S. 'Mabahiss' where deep-sea conoideans were collected. 



No. 


Position 






Area 


Date 


Depth, m 


Gear 


26 


12°29'30"N, 


50°51'30' 


E 


Gulf of Aden 


11.10.1933 


2312 


AT 


33 


13°41'00"N, 
13°40'00"N, 


48°17'00' 
48°18'00' 


Eto 
E 


Gulf of Aden 


15.10.1933 


1295 


AT 


34 


13°05'36"N, 


46°24'42' 


E 


Gulf of Aden 


16.10.1933 


1022 


AT 


35 


13°14'24"N, 
13°13'24"N, 


46° 14' 12' 
46°10'00' 


Eto 
E 


Gulf of Aden 


16.10.1933 


457-549 


OT 


42 


17°26'00"N, 


55°49'00' 


E 


Hadramaut 


27.10.1933 


1415 


TD 


62 


22°53'30" N, 
22°56'30" N, 


64°56'10' 
64°56'30' 


Eto 
E 


Gulf of Oman 


18.11.1933 


1893 


AT 


106 


05°38'54" S, 
05°40'18"S, 


39°15'42' 
39°17'36' 


Eto 
E 


Zanzibar 


12.01.1934 


183-194 


AT 


107 


05°15'30"S, 
05°17'14"S, 


39°33'00' 
39°32'48' 


Eto 
E 


Zanzibar 


12.01.1934 


42 1-457 


AT 


118 


04°05'54" S, 
04°17'00"S, 


41°10'12' 

41 11'48' 


Eto 
E 


Zanzibar 


17.01.1934 


1789 


AT 


119 


06°29'24" S, 
06°32'00" S, 


39049.54. 
39°53'30' 


Eto 
E 


Zanzibar 


19.01.1934 


1207-1463 


AT 


122 


05°21'24"S, 
05°22'36" S, 


39°23'00' 
39°22'18' 


Eto 
E 


Zanzibar 


22.01.1934 


732 


OT 


143 


05°15'48"S, 
05°13'42"S, 


73°22'48' 
73°23'36' 


Eto 
E 


Maldive Is. 


30.03.1934 


797 


AT 


145 


04°58'42" S, 


73°16'24' 


E 


Maldive Is. 


02.04.1932 


494 


AT 


158 


04°42'30" S, 
04°36'48" S, 


72°42'30' 
72°48'54' 


Eto 
E 


Maldive Is. 


07.04.1934 


786-1170 


AT 


176 


12°04'06"N, 


50°38'36' 


E 


Gulf of Aden 


02.05.1934 


665-732 


AT 


180 


12°03'24"N, 


50°40'12' 


E 


Gulf of Aden 


02.05.1934 


397 


G 


184 


14°36'06"N, 
14°38'42"N, 


51°00'18' 
50°57'42' 


Eto 
E 


Gulf of Aden 


04.05.1934 


1270 


AT 


185 


13°48'06"N, 


49°16'48' 


Eto 


Gulf of Aden 


05.05.1934 


2000 


AT 




13°48'36"N, 


49°16'24E 










188 


13°43'18"N, 


47°56'54' 


Eto 


Gulf of Aden 


06 05.1934 


528 


AT 




13°46'00"N, 


47°50'42 E 










193 


13°06'12"N, 


46°24'30' 


Eto 


Gulf of Aden 


07.05.1934 


1061-1080 


AT 




13°03'00"N, 


46°21'42"E 











AT - Agassiz trawl, OT - otter trawl, TD - triangular dredge, G - grab. 

The new species resembles Drillia tasconium Melvill & 
Standen, 1901 from the Persian Gulf but differs in the high spire, 
larger and more solid shell, and absence of spiral sulci which 
deeply furrow the subsutural area in D. tasconium. 

Distribution. Gulf of Aden, 655-732 m. 



Genus HORAICLAVUS Oyama in Taki & Oyama, 1954 

Type species: Mangelia splendida A.Adams, 1867 (original 
designation) 

Horaiclavus splendidus (A. Adams, 1867) 

Figs 20 & 21 

Mangelia splendida. A.Adams, 1867, p. 309, pi. 19, fig. 24. 
Horaiclavus splendidus (A.Adams) -Shuto, 1965, p. 154-155, pi. 

29, figs 13-15, text-figs 3, 5; Powell, 1966, p. 142, pi. 23, fig. 13; 

Habe, 1970, p. 120, pi. 38, fig. 13; Kuroda et al, 1971, p. 

212-213, pi. 55, fig. 4; Shuto, 1975, p. 166, pi. 6, fig. 17. 

Type locality. Goto Islands, Japan. 

Material, stn 176, 3 shells; 188, 1 shell. 

All the JME shells, unlike the specimens illustrated elsewhere, 
are uniformly coloured; the colour has probably faded since the 



time of collection. The protoconch consists of about 2.5 whorls, 
with very small and adpressed initial volution. This is more than 
in the holotype (Shuto, 1 975, pi. 6, fig. 1 7; though it is impossible 
to evaluate the exact number of volutions from the lateral side of 
the illustrated shell) and in Pleistocene shells (1 2/3 whorls, 
Shuto, 1965). 

The JME shells are more similar to Japanese shells than to the 
geographically closer H. madurensis (Schepman, 1913). The 
latter species has somewhat broader and much smaller shell 
(holotype is 14.8 mm in height at 8 teleoconch whorls, according 
to Shuto, 1970, vs. 27.8-32.8 mm at 8-9 whorls in the JME 
shells) with shorter canal and weaker ribs. However in fact these 
species differ only slightly from each other, and H. madurensis 
seems to have no more than subspecific status. 

The familial position of Horaiclavus is still uncertain. Its 
radula was never figured, though Oyama (Taki & Oyama, 1954) 
mentioned that it is similar to that of Comitas and Inquisitor. 
Shuto (1983) described the radula of Horaiclavus as 'true 
toxoglossate according to Kuroda, Habe and Oyama (1971, p. 
327)'. However, the cited page contains no information about 
this genus. Until the examination of the radula, I have to follov 
Shuto (1975, 1983) in assigning Horaiclavus to Drilliidae. 

Distribution. Japan, 50-200 m, and Gulf of Aden, 528-732 
m. The present record is a considerable geographic as well as 
bathymetric range extension for the species. 



DEEP-SEA CONOIDEAN GASTROPODS 










Figs 1-5 Radulae. 1 - Splendrillia zanzibarica Sysoev, new species, holotype; 2 - Inquisitor adenicus Sysoev, new species, paratype, stn 33, H = 34.4 
mm; 3 - /. indistinctus Sysoev, new species, paratype, stn 145, H = 27.4 mm; 4 - Typhlomangelia adenica Sysoev, new species, paratype, stn 1 85, H = 
7.7 mm; 5 - Borsonia (Cordieria) symbiophora Sysoev, new species, paratype, stn 1 85, H = 21 .8 mm. Scale-line = 0. 1 mm. 



A.V. SYSOEV 




Figs 6-13 Radulae (6-9) and opercula (1 0-1 3). 6 - Typhlomangelia maldivica Sysoev, new species, paratype, stn 143, H = 27.8 mm; 7 - Bathytoma 
(Parabathytoma) oldhami (E. A. Smith, 1899), stn 145, H = 41. mm; 8- fi (P.) regnans MdviW, 1918, stn 34, H = 26.6 mm; 9- B. ( P. ) fissa (von 
Martens, 1 901 ), stn 1 76, H = 35.3 mm; 10 - Splendrillia zanzibarica Sysoev, new species, holotype; 1 1 - Inquisitor indistinctus Sysoev, new species, 
paratype, stn 145, H = 27.4 mm; 12, 13-Borsonia (Cordieria) symbiophora Sysoev, new species, paratypes, stn 185, H = 21.8 mm (12) and stn 26, H 
= 20. 1 mm ( 1 3). Scale-lines 0. 1 mm (6-9) and 1 mm (10-13). 



DEEP-SEA CONOIDEAN GASTROPODS 





14 



15 





17 



Figs 14-17 Opercula. 14 - Typhlomangelia adenka Sysoev, new species, paratype, stn 1 85, H = 7.7 mm; 15 T. maldivica Sysoev, new species, para- 
type, stn 143, H = 27.8mm; \6- Bathytoma ( Parabathytoma) regnans Me\\i\\, 1918, stn 34, H = 26.6mm; 17 B. (R)fissa (von Martens, 1901), 
stn 176, H = 35.3 mm. Scale-line = 0.1 mm. 



Genus SPLENDRILLIA Hedley, 1922 

Type species: Drillia woodsi Beddome, 1883 (original 
designation) 

Splendrillia zanzibarica Sysoev, new species 

Figs 1, 10 & 22 

Material, stn 119, 1 specimen (holotype, No. 1993089) and 1 
shell (paratype, No. 1993090). 

Description of holotype. The shell is of medium size for the 
genus, slender, angularly claviform, grayish-white with dull 
surface, rather thin, consisting of 5.5 remained whorls. The 
protoconch is missing; the bluntly closed beginning of initial 
teleoconch whorls is seen in the shell apex. The whorls are angled 
slightly above the periphery, concave above the angulation and 
almost flat below it. Sutures are clear, shallow, slightly wavy. The 
spire is high, occupying about 0.4 of the shell height. Axial 
sculpture consists of strong oblique folds, rapidly disappearing 
on subsutural slope and forming pointed tubercles at the whorl 
periphery. Intervals between folds are narrower than the folds 
themselves. The folds reach the lower part of the shell base but 
weaken greatly on the last third of the body whorl. There are 
about 15 folds on the body whorl and 12 on the penultimate. 
Spiral sculpture is absent except for very indistinct striation seen 
only on the shell base near aperture. The aperture is oval and 
gradually narrows towards the canal. The inner lip is covered by 
a longitudinally rugose callus. The anal sinus is rather shallow, 
broadly open, with the edge somewhat spout-like. The canal is 
moderately long and wide. H = 19.3, Hb = 1 1.7, Ha = 9.3, D = 
8.1 mm. 

The operculum is oblanceolate, with a terminal nucleus. 
Radula is typical for the genus, with small central and comb-like 
lateral teeth. Each marginal tooth is accompanied by a 
transparent, usually more or less triangular plate at its base 
(better seen in detached teeth) which is probably a part of 



radular membrane serving as a tooth ligament. Mean length of 
marginal teeth is 0.26 mm. 

The shell of the paratype (H = 19.4 mm) is badly worn and 
does not differ from the holotype in essential characters. 

The species is characterized by strongly angled whorls, 
oblique axials, and, especially, long canal. It resembles the East 
African Drillia indra Thiele, 1925 while differing in having 
almost 3 times larger shell with long axials and without a distinct 
angulation at the shell base. 

Distribution. Zanzibar, 1207-1463 m. 



Family TURRIDAE H. & A.Adams, 1853 

Subfamily CLAVATULINAE 

Genus TURRICULA Schumacher, 1817 

Type species: Turricula flammea Schumacher, 1817 (monotypy) 
(= Murex tornatus Dillwyn, 1817) 

Turricula new species 

Figs 26 & 30 

Material, stn 119, 1 shell. 

A single young shell (H = 21 .0 mm) apparently represents a new 
species. It resembles most closely the shallow-water species 
Turricula catena (Reeve, 1843) distributed in the Red Sea and 
Persian Gulf, but clearly differs in the absence of the subsutural 
fold sculptured with axial ribs and in the absence of spiral 
sculpture on the shell base. In general shell outline it also 
resembles Surcula fulminata var. gloriosa Melvill, 1917, which 
Powell (1969) dismissed as an abnormal variant of Turricula 
tornata fulminata (Kiener, 1839^10), but is distinguished in its 
heavy nodulose peripheral carina and almost complete absence 
of spiral ribs except for those on the canal. However it seems 



A.V. SYSOEV 




DEEP-SEA CONOIDEAN GASTROPODS 



reasonable to refrain from formal description of a new species 
based on a single young shell. 

Distribution. Zanzibar, 1207-1463 m. 



Subfamily COCHLESPIRINAE Powell, 1942 
Genus COCHLESPIRA Conrad, 1865 

Type species: Pleurotoma cristata Conrad, 1847 (monotypy) 

Cochlespira travancorica (E.A.Smith, 1896) 

Fig. 25 

Pleurotoma (Ancistrosyrinx) travancorica E.A.Smith, 1896, p. 

368; 1906, p. 163; Alcock & Anderson, 1898, pi. 7, figs 1,1a. 
Pleurotoma (Ancistrosyrinx) travancorica var. granulata 

E.A.Smith, 1904, p. 459. 
Ancistrosyrinx travancorica var. granulata (E.A.Smith) - 

Schepman, 1913, p. 420. 
Cochlespira travancorica travancorica (E.A.Smith) & C. 

travancorica travancorica forma granulata (E.A.Smith) - 

Powell, 1969, p. 396-397, pi. 307. 

Type locality, off Travancore (India), 460 fms {travancorica), 
'Investigator', stn 229, 360 fms (var. granulata). 

Material, stn 107, 1 shell. 

The JME specimen was determined and described by Powell 
( 1 969: 396-397) as C. travancorica travancorica forma granulata. 
However this specimen differs from the latter variety (in 
accordance with both original and Schepman's (1913: 420) 
descriptions) in the absence of granules on the spiral ribs of the 
shell base and subsutural slope. The only character in common 
with the variety granulata, is the presence of weak spiral ribs on 
the subsutural slope. At the same time, these differences are 
taxonomically insignificant since the variety granulata, as it was 
mentioned by Powell, apparently does not warrant even a 
subspecific status and represents a form of intraspecific 
variability (this, in particular, makes superfluous the recognition 
of the nominotypical subspecies by Powell ( 1 969: 396)). 

Distribution. East Africa, India, Indonesia, 338-743 m. 

Cochlespira zanzibarica Sysoev, new species 

Figs 23 & 24 

Material, stn 119, 1 shell (holotype, No. 1993091). 

Description. The shell is small, light-brown, with glossy 
surface, rather solid, fusiform, consisting of 6.5 preserved 
whorls. The protoconch is missing and the upper whorls are 
heavily eroded. The whorls are angled below the periphery and 
concave upper and below the angulation; the position of the 
angulation on the spire whorls shifts upward towards the body 
whorl. The sutures are straight and very shallowly impressed. 
The sculpture consists only of a strong median keel with 
rounded, pointed tubercles (16 on the body and penultimate 



whorls) and a low ridge on the upper part of the shell base. The 
ridge is hard to trace above the suture on last spire whorls. The 
growth lines are very thin. The shell base is angled in its upper 
part, where the ridge is situated, and slightly and evenly concave 
below the angulation; it smoothly passes into the canal. The 
aperture is narrow, with the inner lip weakly and evenly curved. 
The anal sinus is broad and moderately deep, with the apex 
situated in the middle of subsutural slope. The canal is straight 
and rather short (the end is apparently slightly broken). H = 
16.9, Hb = 10.2, Ha = 7.6, D = 6.9 mm. 

The new species obviously belongs to the 'semiplana group' 
sensu Powell, 1969 which includes two fossil and two Recent 
deep-water species and is characterized by the presence of a 
strong basal keel. The new species differs from all known species 
of the group by the complete absence of spiral sculpture on the 
shell base, except for low ridge and in its more stout shell with 
rather short canal. It is also similar to species of the genus 
Chesasyrinx Petuch, 1988 known from Miocene of Maryland, 
USA. Although Petuch (1988: p. 38-39) did not compare 
Chesasyrinx with Cochlespira in the original description, the 
striking similarity of shells of Chesasyrinx and 'semiplana 
group' of Cochlespira may be reason for synonymizing these 
genera. 

Distribution. Zanzibar area, 1207-1463 m. 



Genus COMITAS Finlay, 1926 

Type species: Drillia fusiformis Hutton, 1877 (= Surcula huttoni 
Suter, 1914) (original designation) 

Comitas subsuturalis (von Martens, 1902) 

Figs 31^10 

Pleurotoma ( Brachytoma) subsuturalis von Martens, 1902, p. 

239. 
Brachytoma subsuturalis (von Martens) — von Martens, 1903 

[1904], p. 85, pi. 1, fig. 7. 
Comitas subsuturalis (von Martens) — Powell, 1969, p. 285, pi. 

226, figs 3^1. 

Type locality. 'Valdivia', stn 256, off Somali, 1 134 m. 
Material, stn 33, 3 specimens and 4 shells 

and 8 shells - stn 118 1 snerimen anH 1 



and 8 shells; stn 118, 1 specimen am. 
specimens; stn 145, 2 specimens; stn 184, 7 
stn 193, 3 shells. 



„, stn 34, 1 specimen 

specimen and 1 shell; stn 143, 9 

7 specimens and 1 shell; 



Examination of the large series of JME specimens revealed very 
a high variability of C. subsuturalis in sculpture and shell 
proportions. Some specimens are very similar to the original 
illustration of von Martens (e.g. Figs 37, 38) whilst others, often 
from the same station, may differ in narrower (or, conversely, 
broader) shells with more or less high position of peripheral 
keel, variously differentiated spiral ribs on the shell base, and 
more or less prominent and numerous tubercles on subsutural 
fold. The largest JME shell is 32.4 mm in height and 12.0 mm in 
width. 



Figs 18-30 Clavusidae, Clavatulinae and Cochlespirinae. 18, 19 - Drillia altispira Sysoev, new species, holotype; 20, 21 - Horaiclavus splendidus (A. 
Adams, 1 867), stn 1 88 (20) and 1 76 (21 ), H = 32.4 (20) and 32.8 (21 ) mm; 22 - Splendrillia zanzibarica Sysoev, new species, holotype; 23, 24 - Coch- 
lespira zanzibarica Sysoev, new species, holotype; 25 - Cochlespira travancorica (E.A. Smith, 1 896), stn 107, H = 19. 1 mm; 26, 30 - Turricula new 
species, stn 119, H = 21.0 mm; 27- Comitas elegans Sysoev, new species, holotype; 28 - Leucosyrinx claviforma (Kosuge, 1992), stn 158, H = 28.0 
mm; 29 - Comitas curviplicata Sysoev, new species, holotype. 



A.V. SYSOEV 



Powell (1969, p. 285) mentioned 'a related new species from 
the Gulf of Aden in 1270 metres', i.e. from stn 184, but did not 
give a formal description nor reasons for this. However, extreme 
variants of C. subsuturalis from stn 184 are connected by 
intermediate forms and can be therefore identified as that 
species. 

Specimens from two stations off Maldive Islands comprise a 
distinct group differing from East African shells in smaller size 
(15.5-21.0 mm at 6-7 teleoconch whorls vs. 21.8-28.7 in typical 
C. subsuturalis) and more slender shell proportions (H/D ratio is 
2.82-3.12, mean 2.95 (n = 1 1) vs. 2. 10-2.94 (2.10-2.70 in 19 out 
of 20 shells measured), mean 2.34). These differences are 
probably connected with geographical isolation of the Maldive 
Islands resulting in formation of morphologically isolated 
population of the species. If the above-mentioned metric 
differences will be confirmed in additional samples, the 
population of C. subsuturalis from Maldive Islands should be 
considered as a distinct subspecies. This population represents a 
transition (both geographical and conchological) to C. exstructa 
von Martens, 1903, described from Nicobar Islands. The latter 
species is distinguished only by an even narrower shell (H/D 
ratio is 3.43 in the holotype) with longer axial folds (as far as it 
can be judged from von Martens' figure). Examination of type 
material may however reveal that C. exstructa is a synonym of C. 
subsuturalis. A similar statement is probably true for C. 
obtusigemmata Schepman, 1913, which does not differ from C. 
subsuturalis in essential conchological characters. 

Distribution. East Africa from the Gulf of Aden to Zanzibar, 
and Maldive Islands, 494-1789 m. 



Comitas erica (Thiele, 1925) 

Fig. 41 

Leucosyrinx erica Thiele, 1925, p. 236, pi. 36(24), fig. 25. 
Comitas erica (Thiele) — Powell, 1969, p. 284, p. 226, fig. 2. 

Type locality. 'Valdivia', stn 191, off Siberut Id. (Sumatra), 
750 m. 

Material, stn 143, 2 specimens. Also mentioned by Powell 
(1969) from stn 108 (Zanzibar area, SE of Pemba Island, 786 m). 

Both specimens are very similar to the original figure and the 
species variability thus seems to be rather low. One of JME 
specimens is larger than Thiele's holotype (20.9 vs. 1 6. 5 mm), but 
has the appearance of an immature shell. The species was found 
in the sample also containing C. subsuturalis, but it can be easily 
distinguished from the latter, by the grayish-white shell with 
weaker subsutural tubercles, closer-spaced spiral ribs, two of 
which on the upper shell base are more prominent, and larger 
protoconch (0.95 mm in diameter vs. 0.7-0.8 mm in C. 
subsuturalis from the same sample). 

Distribution. Sumatra, Zanzibar and Maldive Islands, 
750-797 m. 



Comitas pauper a (Watson, 1881) 

Figs 42^8 

Pleurotoma (Drillia) paupera Watson, 1 88 1 , p. 41 1 . 
Pleurotoma (Typhlomangelia) paupera Watson - Watson, 

p. 317-319, pi. 25, fig. 3. 
Turricula paupera Watson -Powell, 1969, p. 244, pi. 202. 



886, 



Type locality. 'Challenger', stn 191, off the Arrou Islands 
(Arafura Sea, Indonesia), 800 fms. 

Material, stn 62, 2 specimens; stn 185, 3 specimens and 4 
shells. 

Proper determination of this species is rather intricate due to the 
very heterogeneous type material. Powell (1969) erroneously 
described Watson's syntypes as consisting of two shells with 
rounded lower whorls and obsolescent sculpture and one shell 
angulate with strong axially costae. He illustrated the latter 
specimen and designated it as the holotype (correctly named 
lectotype in the 'Measurement' and 'Types' paragraphs). 
Actually, one of the two paralectotypes (H = 33.3 mm) has a 
rounded body whorl with obsolete axials while the axial 
sculpture on spire whorls is almost the same as in the lectotype 
(Figs 42 & 43). The second paralectotype (Fig. 44) is represented 
by a small (H = 18.5 mm) and quite dissimilar shell, 
characterized by strongly angled whorls with axial sculpture 
consisting of strong and rounded tubercles at the place of whorl 
angulation. This specimen is so distinct that one could easily 
assign it to a separate species if it was not found in the same 
sample. 

Two at first glance rather different shells from the JME stn 62 
fit however in the range of variability described above. The larger 
shell (H = 23.4 mm, Fig. 45) is comparable to the lectotype, but 
has less convex whorls with a less pronounced subsutural slope. 
The smaller shell (H = 17.5 mm, Fig. 46) is very similar to the 
smaller paralectotype, but has even stronger peripheral nodules. 
Small specimens (14—15 mm in height) from stn 185 (Figs 47 & 
48) have shells and sculpture intermediate between the extreme 
variants. In some of the latter specimens, the spiral ribs are more 
widely spaced on the shell base and reduced in number on the 
subsutural slope. 

Thus, C. paupera appears to be very variable species with 
extreme variants being quite dissimilar to each other but 
connected by intermediate forms. 

The species was included by Powell (1969) into the genus 
Turricula Schumacher, 1817. However its operculum has a 
terminal nucleus characteristic of the genus Comitas. 

Surcula suratensis Thiele, 1925 (= Surcula coreanica of von 
Martens, 1903, not of Adams & Reeve, 1850) is similar to some 
forms of C. paupera and after examination of type material it 
may appear to be a synonym of the latter. 

Distribution. Indonesia, Gulf of Aden, and Gulf of Oman, 
1463-2000 m. 



Figs 31^41 Comitas subsuturalis (von Martens, 1 902) (31^10) and C. erica (Thiele, 1 925) (41 ). 31, 32 - stn 1 84, H = 37.3 mm; 33 - stn 1 84, H = 25.5 
mm; 34 -stn 184, H = 24.0 mm; 35 -stn 33, H = 21. 0mm; 36 -stn 184, H = 28.6 mm; 37 -stn 34, H = 27.0 mm; 38 -stn 184, H = 21. 7 mm; 39 -stn 
143, H = 21 .0 mm; 40 - stn 145, H = 26.2 mm; 41 - stn 143, H = 20.9 mm. 



DEEP-SEA CONOIDEAN GASTROPODS 




10 



A.V. SYSOEV 




Figs 42-48 Comitas paupera (Watson, 1 881 ). 41 - lectotype, H = 40.0 mm; 43, 44 - paralectotypes, H = 33.3 (43) and 1 8.5 (44) mm, BM(NH) 
1887.2.9.1009-ll;45,46-stn62,H = 23.4(45)and 17.5 (46) mm; 47,48 -stn 185, H = 14.0 (47) and 14.9(48) mm. 



Comitas cwviplicata Sysoev, new species 

Fig. 29 

Material, stn 184, 2 specimens (holotype No. 1993092 and 
paratypeNo. 1993093). 

Description of holotype. The shell medium size, fusiform, 
rather thin, covered with light-brown periostracum, consists of 7 
whorls. The protoconch is missing. The whorls are angled at the 
periphery or below it (on upper whorls). The subsutural slope is 
broad and weakly concave. Sutures are slightly wavy, shallow. 
Axial folds are low, reversed-sigmoid, they form strong rounded 
nodules below the subsutural slope and then rapidly weaken and 
become narrowly crested, but extend to the lower suture on the 
spire whorl and to lower part of the shell base. There are 1 5 axial 
folds on the body whorl and 13 on the penultimate. The 
subsutural slope is devoid of axial sculpture. Growth lines are 
mostly indistinct except those on the subsutural slope, but some 
are rough and raised. Spiral ribs are unequal in width and 



prominence, generally they are most closely spaced in intervals 
between peripheral nodules and further apart on the lower shell 
base and the canal. Two ribs on the upper shell base are raised 
and most prominent. The subsutural slope is covered with 
subobsolete, closely set ribs which are obsolete on some parts of 
the shell; there are also two shallow grooves in the middle of 
subsutural slope. The body whorl is large, occupying about 0.66 
of the shell height. The shell base is almost flat, evenly curved in 
transition to the canal. The aperture is oval, with the inner lip 
evenly curved and covered by a wide callus which bears oblique 
folds being the continuation of spiral ribs. The anal sinus is 
moderately deep, broadly V-shaped, with its apex being in the 
middle of subsutural slope. The canal is moderately long, 
slightly turned to left. The operculum is large, leaf-shaped, with 
terminal nucleus. H = 32.4, Hb = 21 .6, Ha = 17.6, D = 13.0 mm. 

The paratype (H = 29.2, D = 11.8 mm) is similar to the 
holotype, but has rather worn surface. 

The new species is similar to C. arcana (E.A.Smith, 1 899) from 
Andaman Islands and Southern India (338-658 m) but differs in 



DEEP-SEA CONOIDEAN GASTROPODS 



11 



having a slightly larger shell with shorter and broader canal, 
shorter spire (its height is less than that of aperture plus canal), 
and long axials with characteristic reversed-sigmoid curvature 
and less strong and more numerous nodules. 

Distribution. Gulf of Aden, 1270 m. 

Comitas elegans Sysoev, new species 

Fig. 27 

Material, stn 176, 1 shell (holotype, No. 1993094). 

Description. The shell is medium size, fusiform, slender, with 
a high turreted spire, solid, white under a grayish-brown 
periostracum, and consists of 8 whorls. The protoconch is 
missing. Whorls are roundly angled at the shoulder, with 
conspicuous subsutural fold, slightly concave subsutural slope, 
and almost vertical lateral sides. Sutures are shallow, wavy, and 
slightly channelled. Axial folds (12 on the body whorl and 1 1 on 
the penultimate one) are strong, oblique, broad, and rounded. 
They rapidly disappear on subsutural slope and slightly weaken 
towards the lower suture. On the body whorl, the folds are 
present on the whorl periphery and fade on the upper part of the 
shell base. Intervals between folds are narrower than the folds 
themselves. Spiral ribs are low, broad, rounded and divided by 
narrow interstices in the upper part of whorls below the 
subsutural slope. Towards the lower suture and on the shell base 
they become narrower, thread-like, and more widely spaced. 
Subsutural slope is smooth except for indistinct spiral grooves in 
the middle. Growth lines are numerous, thin, prominent on the 
subsutural slope. The shell base is almost flat; weakly curving, it 
passes smoothly into a long and straight canal. The aperture is 
narrow, elongate-oval and not differentiated from the canal. The 
inner lip is almost straight along most of its length, covered by 
wide but thin callus. The anal sinus, judging from growth lines, is 
rather deep, V-shaped, with the apex situated just below the 
middle of subsutural slope. H = 38.3, Hb = 24.7, Ha = 19.0, D = 
11.6 mm. 

The new species is very similar to Drillia investigatoris 
E.A.Smith, 1899 and differs in having a much smaller shell (65 
mm in the unique holotype of D. investigatoris) with subsutural 
slope devoid of spiral sculpture. 

Distribution. Gulf of Aden, 655-732 m. 

Comitas sp. 

Material, stn 184, 1 shell. 

A single broken shell (H = 24.4, upper spire whorls missing) is 
rather similar to C. trailli (Hutton, 1 873) from New Zealand, but 
the worn condition of the shell hampers its proper identification. 



Genus LEUCOSYRINX Da\\, 1889 

Type species. Pleurotoma verrilli Dall, 1881 (original 
designation) 

Leucosyrinx claviforma (Kosuge, 1992) 

Fig. 28 

Comitas claviforma Kosuge, 1992, p. 165-166, pi. 58, figs 7-8. 
Type locality, off Port Hedland, Western Australia, 376 m. 
Material, stn 158, 1 shell. 



The JME shell differs from the unique holotype in having 
slightly less angled whorls, lower spire (though the shell is 
smaller: H = 28.0 mm vs. 38.1 mm in the holotype), and reddish 
brown colour (ashy white in the holotype). However, all other 
conchological characters agree with original description and 
illustrations. 

The species was described as Comitas, but the narrow, turreted 
shell with angled whorls and the anal sinus scars indicating a 
deep sinus with greatly projected outer lip are more typical of the 
genus Leucosyrinx. 

Distribution. Western Australia and Maldive Islands, 
376-1 170 m. 



Subfamily CRASSISPIRINAE Morrison, 1966 
Genus INQUISITOR Hedley, 1918 

Type species: Pleurotoma sterrha Watson, 1881 (original 
designation) 

Inquisitor nodicostatus Kilburn, 1988 

Fig. 49 

Crassispira aesopus (non Schepman, 1913) - Kilburn, 1973, p. 

572, fig. 13a. 
Inquisitor nodicostatus Kilburn, 1988, p. 259-261, figs 36, 42, 

213-214. 

TYPE LOCALITY. Natal, South Africa (29°43'S, 31°05'E), 
164-169fms. 

Material, stn 106, 1 shell. 

Distribution. South Africa and Zanzibar, 183-310 m. 

Inquisitor adenicus Sysoev, new species 

Figs 2 & 50-53 

Material, stn 33, 1 paratype No. 1993096/1; stn 34, 7 
paratypes No. 1993096/2-8; stn 193, holotype No. 1993095 and 
4 paratypes No. 1993096/9-12. 

Description of holotype. The shell is medium sized for the 
genus, claviform, thin but solid, with a high spire, covered with 
olivaceous-brown periostracum, and consisting of 8 whorls. The 
protoconch is missing and the upper whorls are eroded. Whorls 
are obtusely angled at the shoulder and moderately convex. 
Subsutural slope is broad and concave and the subsutural fold is 
weak and indistinct. Sutures are shallow and wavy. Axial folds 
are strong and rounded, slightly oblique, with interstices 
narrower than folds. Folds extend from the lower suture to the 
lower part of subsutural slope where they rapidly disappear. 
There are ten folds on each of two last whorls. Growth lines are 
thin, prominent on the subsutural slope. Spiral cords are strong, 
rounded, almost equally developed on axial folds and in 
interstices; they are much narrower than the interspaces between 
them. The cords are absent on subsutural slope and become 
narrower, closer and much weaker on lower part of the shell base 
towards the canal end. There are four cords on the spire whorls 
(five on the penultimate one, the lowest cord submargins the 
suture), five on the body whorl periphery, and, below a wider 
interval, about 12 on the shell base and canal. The aperture is 
elongate-oval, rather narrow, becoming slightly narrower as it 
passing into the canal. Outer lip with thin edge; inner lip 



12 



A.V. SYSOEV 



concave, covered with wide, longitudinally rugose callus, which 
does not form a pad at the sinus entrance. The anal sinus is deep, 
narrows markedly towards the apex, U-shaped (type (c) of 
Kilburn, 1988), and its deepest point is situated slightly below 
the middle of subsutural slope. The canal is moderately long and 
straight. H = 33.0, Hb = 18.4, Ha = 13.8, D = 10.3 mm. 

The paratypes vary slightly in the character of spiral 
sculpture: the cords may be unevenly spaced, rarely with thinner 
additional cords in some intervals. The subsutural fold is 
variously developed, sometimes it is distinct. The canal is 
narrower and sometimes curved in smaller shells and broader 
and therefore visually shorter in larger paratypes. The largest 
paratype is 34.3 mm in height. 

In the only paratype with an intact protoconch, it consists of 
1.5 rapidly increasing globose whorls ( 1 .0 mm in diameter) with 
a smooth surface. An operculum was not found; it might be lost 
in the dried animal. The radula (Fig. 2) is typical of the genus, 
teeth with tapering distal end of the shaft, poorly differentiated 
cutting edge, and large and broad accessory limb. The tooth 
length is 0.1 7 mm (in paratype from stn 33, H = 34.3 mm). 

The new species resembles Funa laterculoides (Barnard, 1958) 
in general outlines but clearly differs in its protoconch, colour, 
radula, and details of sculpture. 

Distribution. Gulf of Aden, 1022-1295 m. 

Inquisitor indistinctus Sysoev, new species 

Figs 3, 1 1 & 54-55 

Material, stn 145, 3 specimens and 1 shell (holotype No. 
1993097 and 3 paratypes No. 1993098). 

Description of holotype. The shell is medium size, 
claviform, slender, with rather high spire, thin but solid, covered 
with solid olivaceous periostracum, and consists of protoconch 
and 9 teleoconch whorls. The protoconch is small (0.95 mm in 
diameter) consisting of 1.5 smooth glossy whorls. Definitive 
whorls weakly convex, with a slight angulation at the shoulder. 
Subsutural slope concave except for weak subsutural fold. 
Sutures straight, moderately deep, become slightly channelled 
on the body whorl. The axial sculpture is represented by strong 
oblique rounded folds, gradually disappearing on subsutural 
slope and reaching the lower suture or, on the body whorl, the 
shell base. The folds tend to weaken on the last half of the body 
whorl. There are 14 folds on the body and penultimate whorls. 
Spiral cords override the axial folds, they are low, wide and 
rounded, the intervals are approximately equal to cords in width 
or somewhat wider. There are 16 cords on the body whorl and 
7-8 on the penultimate. The shell base is weakly convex, and 
smoothly passes into the canal. The aperture is rather narrow, 
elongate-oval, and not differentiated from the canal. The latter is 
short and wide, somewhat expanded and shallowly notched at 
the end. The anal sinus is moderately deep, rounded, 
symmetrical, and occupies the entire subsutural slope. The inner 
lip is covered with thick white callus with a free edge in the lower 
part. Columella almost straight. H = 31.3, Hb = 16.5, Ha = 12.4, 
D = 8.3mm. 

The paratypes are smaller (H no more than 27.4 mm) and vary 
slightly in the prominence of the axial and spiral sculpture 



including the subsutural fold. In one paratype, there is a weak 
but distinct callus pad at the entrance to anal sinus. The 
operculum is oblanceolate, with terminal nucleus. Marginal 
teeth of the radula possess broad leaf-shaped accessory limb, 
which does not reach the distal end of the shaft. The mean tooth 
length is 0. 14 mm (in paratype with H = 27.4 mm). 

Distribution. Maldive Islands, 494 m. 

Inquisitor stenos Sysoev, new species 

Figs 56 & 57 

Material, stn 176, 1 shell (holotype, No. 1993099). 

Description. The shell is rather small, slender, with high spire 
comprising about 0.5 of the shell height, yellowish-white, and 
consisting of 9 whorls. The protoconch is missing. Whorls are 
strongly angled at the periphery, and the whorl profile is very 
weakly concave above the angulation and almost flat below it. 
There is a narrow and weak subsutural fold. Sutures are straight 
and moderately deep. The axial sculpture consists of slightly 
oblique and widely spaced folds (9 on each of two last whorls) 
forming longitudinally elongated and sometimes pointed strong 
tubercles in the middle of the whorl. On early spire whorls, the 
folds are obsolete but visible on the lower part of subsutural 
slope and reach the lower suture. Towards the body whorl, they 
become obsolete near the lower suture and over most part of 
subsutural slope, but again become longer and extend over entire 
shell base in the last half of the body whorl. The last fold situated 
behind the aperture edge is much wider and stronger than other. 
Spiral sculpture consists of indistinct, rather broad ribs which 
are obsolete or subobsolete in interstices between axial folds and 
absent on the subsutural slope. The aperture is narrow, 
elongate-oval, with a thick labial callus and straight columella. 
The labrum has a thin edge and low and narrow fold-like varix 
behind the edge. The anal sinus is moderately deep, U-shaped, 
with slightly constricted entrance (type (b) of Kilburn, 1988). 
There is a moderately large, pointed, and outwardly projecting 
parietal tubercle. H = 21.3, Hb = 10.7, Ha = 18.4, D = 5.7 mm. 

The new species is distinguished by its small narrow shell with 
high spire, broad subsutural slope, low but strongly tuberculated 
at the periphery axial folds, and obscure spiral ribs. 

Distribution. Gulf of Aden, 655-732 m. 

Inquisitor angustiliratus Sysoev, new species 

Fig. 58 

Material, stn 188, 1 shell (holotype, No. 1993100). 

DESCRIPTION. The shell is moderately large, with a rather high 
spire, strong, yellowish-gray, and consisting of 8 whorls. The 
protoconch is missing. Whorls are moderately convex, slightly 
angled at the shoulder, with indistinct subsutural fold and 
moderately wide (about 1/3 of the whorl height), concave 
subsutural slope. Sutures are wavy, and slightly channelled. 
Axial folds ( 1 2 on the body whorl and 1 1 on the penultimate one) 
are narrow, slightly oblique, with sharpened crests, and long 
(they reach the lower suture and extend over the entire shell 
base). The folds begin in the lower part of the subsutural slope 



Figs 49-62 Crassispirinae. 49 - Inquisitor nodicostatus Kilburn, 1 988, stn 1 06, H = 36.8 mm; 50-53 - /. adenicus Sysoev, new species, holotype (50) 
and paratypes, stn 33 (51) and 34 (52-53), H = 34.3 (51), 30.9 (52) and 26.3 (53) mm; 54, 55 - /. indistinctus Sysoev, new species, holotype (54) and 
paratype, stn 145, H = 26.0 mm (55); 56, 57 - /. stenos Sysoev, new species, holotype; 58 - /. angustilirata Sysoev, new species, holotype; 59, 60 - 
Paradrillia agalma (E.A. Smith, = 1 906), stn 1 76, H = 1 5.3 mm; 61 - P. agalmal, stn 1 80, H = 14.4 mm; 62 - Ceritoturris sp, stn 42, H = 1 1 .8 mm. 



DEEP-SEA CONOIDEAN GASTROPODS 



13 




14 



A.V. SYSOEV 



and are most prominent in the whorl periphery. Spiral ribs are 
strong, cord-like, narrow, widely spaced (but become 
progressively closer to each other and lower on the shell base and 
canal); they override the axial folds and form nodules at 
intersections. Interspaces between ribs are very finely spirally 
striate and sometimes bear a thin thread. The number of ribs 
increases from 3 to 5 in successive spire whorls. The subsutural 
fold is sculptured by two thin spiral riblets. Subsutural slope is 
smooth except for very fine spiral striations and 3-4 thin threads 
in the lower half. Growth lines are numerous, thin, and clear, on 
the body whorl some of them are rough and raised. The shell 
base is almost flat and not differentiated from the canal. The 
aperture seems to be rather wide (the outer lip is broken). The 
inner lip is weakly and evenly curved, and covered by thick 
callus. The parietal callus pad is very weak. The anal sinus 
(judging from its scars) is moderately deep, openly U-shaped 
(type (c) of Kilburn, 1988). The canal is broad and widely open. 
H = 39.5, Hb = 21.4, Ha = 16.5, D = 10.8 mm. 

The new species resembles /. coxi (Angas, 1867) from 
south-eastern Australia in whorl outline and the character of 
sculpture but clearly differs in the shell proportions. It also looks 
somewhat like an extremely stretched out specimen of /. crassa 
(E.A.Smith, U 



Distribution. Gulf of Aden, 528 m. 



Genus PARADRILLIA Makiyama, 1940 

Type species: Drillia dainichiensis Yokoyama, 1923 (original 
designation) 

Paradrillia agalma (E.A.Smith, 1906) 

Figs 59-61 

Pleurotoma (Surcula) agalma E.A.Smith, 1906, p. 162-163; 

Annandale & Stewart, 1909, pi. 21, figs 4, 4a. 
Paradrillia agalma (E.A.Smith) - Powell, 1969, p. 317, pi. 246, 

figs 1,2. 

Type locality. 'Investigator', stn 269, West of Cape Comorin 
(SE India), 464 fms. 

Material, stn 176, 1 shell; stn 180, 2 shells. 

The shell from stn 176 is quite typical. It consists of protoconch 
and 7 teleoconch whorls. The previously undescribed 
protoconch is basically similar to that of P. melvilli figured by 
Powell (1969, pi. 242, fig. 2). It consists of 2.5 whorls with 
smooth glossy surface, the tip is small and papillate, and a thin 
low-set keel is developed on the last whorl. There are 1 7 axial ribs 
on the body whorl and 1 5 on the penultimate one. 

The shells from stn 180 (Fig. 61) were referred to P. agalma 
with some doubts. They differ in more numerous peripheral 
tubercles, weaker axial and spiral sculpture, and in the presence 
of two spiral threads on the subsutural fold and two riblets 
below the periphery on the spire whorls. They seem to be a 
transition to P. melvilli Powell, 1 969 in sculpture, but that species 
is almost half the size, with a proportionally higher spire and 
truncated anterior end. However, the general pattern of 
sculpture, shell outline and the character of protoconch are 
similar in all JME shells and those from stn 1 80 are provisionally 
assigned to P. agalma pending examination of additional 
material. 

Distribution. Ceylon and Gulf of Aden, 655-848 m. 



Genus CERITOTURRIS Da\\, 1924 

Type species: Ceritoturris bittium Dall, 1924 (original 
designation) 

TCeritoturris sp. 

Fig. 62 

Material, stn 42, 1 shell. 

A single heavily damaged shell (H = 1 1.8 mm) from stn 42 can 
possibly be referred to Ceritoturris on the basis of its 
resemblance to both type species and C. nataliae Kilburn, 1988. 
The very bad condition of the shell renders more precise 
identification impossible. 

Distribution. West Arabian Sea, 1415 m. 



Genus PTYCHOBELA Thiele, 1925 

Type species: Clavatula crenularis Lamarck, 1816 (= Murex 
nodulosus Gmelin, 1791) (original designation) 

Ptychobela cf. suturalis (Gray, 1838) 

Figs 79 & 80 

Drillia suturalis Gray, 1838, p. 29. 

Ptychobela suturalis (Gray) - Kilburn, 1989, p. 190, figs 5-6 
(holotype) & 7-8. 

TYPE LOCALITY, unknown. 

Material, stn 35, 1 shell; stn 188, 2 shells. 

These shells are difficult to determine primarily due to the 
existence of numerous species of uncertain status which have 
been described within the genus Drillia and, as far as it can be 
judged from drawings (when present) and rather brief 
descriptions, are similar to the JME material (e.g. Drillia incerta 
E.A.Smith, 1877, D. atkinsoni E.A.Smith, 1877, D. variabilis 
E.A.Smith, 1877, etc.). The question cannot be resolved without 
comparative examination of type material. Nevertheless, these 
shells are quite comparable with the holotype of P. suturalis 
illustrated by Kilburn (1989). The latter species is however 
characterized by usually very short axial folds with strong 
peripheral nodules (but the folds in the holotype are rather long 
- see Kilburn, 1989, fig. 6). The smaller of the JME shells (stn 
188, H = 22.7, D = 7.6 mm, Fig. 80) differs also in complete 
absence of additional spiral threads in the interspaces between 
main cords. The other shell (stn 188, H = 22.9, Hb = 15.6 mm, 
upper spire whorls are broken off and only 3.5 last whorls are 
intact) also lacks additional threads on upper spire whorls, but 
they appear on penultimate whorl and become rather strong on 
the body whorl. The anal sinus is similar in shape to that of P. 
suturalis, but some of the sinus scars indicate that during the 
shell growth the sinus may be quite different: very deep and 
narrow, asymmetrical, with the upper edge of the slit almost 
parallel to the suture. The rather large shell from stn 35 (H = 
3 1 . 1 , D = 11.0 mm, Fig. 79) is characterized by fairly long axials 
reaching the lower suture and a peculiar spiral sculpture of ribs 
with the upper (directed adapically) slope being much steeper 
than the lower. This produces a somewhat tiled pattern and, 
when illuminated from the shell apex, the sculpture seems 
consist of very wide flattened ribs. 



DEEP-SEA CONOIDEAN GASTROPODS 



15 



Distribution. According to Kilburn (1989), P. sutwalis is a 
central West Pacific species (from Taiwan and Singapore to 
Queensland and Western Australia). 

Ptychobela cf. nodulosa (Gmelin, 1791) 

Fig. 84 

Murex nodulosus Gmelin, 1791, p. 3562. 
Clavatula crenularis Lamarck, 1816, p. 9, pi. 440, figs 3a,b. 
Ptychobela nodulosa (Gmelin) - Kilburn, 1989, p. 187-190, figs 
1-2) & 3^1 (neotype). 

Type locality, unknown. 

Material, stn 188, 1 shell. 

The situation with this species is the same as in the preceding 
case. The JME shell resembles rather closely the neotype of P. 
nodulosa designated and illustrated by Kilburn ( 1 989) in the shell 
outline and the character of sculpture. However, it lacks the 
characteristic colour pattern of P. nodulosa being of uniform 
light-brown colour (the shell was dead collected and probably 
faded), and has a slightly higher spire (Hs/H = 0.49 vs. 0.41 in the 
neotype of P. nodulosa) and more convex whorls. A 
characteristic feature of the shell considered is that axial folds 
extend from suture to suture on the uppermost whorls and only 
on 8th teleoconch whorl the typical subsutural slope without 
axial sculpture is developed. 



Subfamily TURRINAE H. & A.Adams, 1953 
Genus GEMMULA Weinkauff, 1875 

Type species: Pleurotoma gemmata Reeve, 1843 (subsequent 
designation Cossmann, 1896) (= Gemmula hindsiana Berry, 
1958) 

Gemmula (Gemmula) vagata (E.A.Smith, 1895) 

Figs 63-65 

Pleurotoma vagata E.A.Smith, 1895, p. 3, pi. 1, fig. 3; 1904, p. 

456;Alcocke/a/., 1907, pi. 14, figs 3, 3a. 
Gemmula vagata (E.A.Smith) - Powell, 1964, p. 258-259, pi. 196, 

fig. 10. 

Type locality. 'Investigator', stn 172, off Trincomalee, 
Ceylon, 200-350 fms. 

Material, stn 176, 8 shells, stn 188, 3 shells. 

The largest shell is 49.8 mm in height (apex slightly broken). 
Examination of a growth series showed that its characteristic 
features (i.e. almost vertical sides of whorls, very strongly 
excavated subsutural fold, and channelled sutures) are 
developed only when the shell reaches a certain size 
(approximately 35 mm in height and more than 10 teleoconch 
whorls). Young shells can be determined only by comparison 
with larger specimens. A peculiar and previously undescribed 
feature of the species is the presence of spiral lirae inside the 
aperture in large individuals. 

Distribution. Gulf of Aden to Andaman Islands, 338-1061 
m. 



Gemmula (Gemmula) amabilis (Jickeli in Weinkauff, 

1875) 

Figs 66, 67 & 71 

Pleurotoma amabilis Jickeli in Weinkauff, 1875, p. 29, pi. 6, figs 

4,6. 
Pleurotoma (Gemmula) amabilis Weinkauff -Sturany, 1903, pi. 

3, figs 3a-c. 
Gemmula amabilis (Weinkauff) - Powell, 1964, p. 261-262, pi. 

200, fig. 1, pi. 201, figs 3-7. 
IGemmula amabilis (Weinkauff) - Kosuge, 1990, p. 153-154, pi. 

55, fig. 13, text-fig. 6; Kosuge, 1992, p. 163, pi. 58, fig. 1, 

text-figs 7, 11,12-14. 

Type locality. Massawa (Ethiopia), Red Sea. 

Material, stn 34, 18 specimens and shells; stn 1 19, 1 shell; stn 
145, 1 specimen and 2 shells; stn 176, 2 shells; stn 185, 1 shell; stn 
188, 2 shells; stn 193, 3 specimens. 

The taxonomy of this species is rather confused. The name 
amabilis had long been unused until Powell ( 1 964) applied it to 
shells from the JME material (the specimen from the JME stn 
176 figured by him (Powell, 1964, p. 200, fig. 1) was not found). 
However, Powell expressed some doubts on the identification of 
JME specimens as G amabilis, having indicated certain 
differences from the sketchy and rather inadequate original 
figures. 

Kilburn (1983) reported that the types of Pleurotoma amabilis 
were probably lost and therefore the name amabilis must remain 
a nomen dubium. He also compared the specimen figured by 
Powell with G. pulchella Shuto, 1961 from the Pliocene of Japan. 
The latter species is characterized by smaller and much more 
numerous gemmules (29 on the body whorl of 10.2 mm holotype 
and up to 34 at the shell height of 26.57 mm - Shuto, 1965). 

Later on, Kosuge (1990, 1992) used the name amabilis for 
North-Western Australian shells and at the same time 
mentioned that Powell's specimen differs from the original 
drawings and, as far as it can be judged from the text (Kosuge, 
1990, p. 154), may not be conspecific with amabilis. The shells 
figured by Kosuge as G amabilis, actually resemble G 
gemmulina (von Martens, 1902) sensu Powell, 1964 more in the 
shell outlines and the character of spiral sculpture of alternating 
primary cords and intermediate threads. The latter species is 
distributed from Taiwan to Indonesia (Powell, 1964), which is 
far closer to North-Western Australia than the Red Sea and the 
Gulf of Aden. 

Despite the obviously ambiguos status of the name amabilis, 
it seems reasonable to conserve its application to the Gulf of 
Aden shells similar to those described by Powell, because no 
other available name for them exists. Perhaps, the examination 
of a large series of the Red Sea Gemmula, especially from the 
type locality of amabilis, would clear up the question of proper 
application of the name. The specimens of G amabilis are 
characterized by a small slender shell (up to 25.9 mm, usually 
22-25 mm) with more or less channelled sutures, a moderately 
developed subsutural fold covered by 1-3 riblets, 2-3 prominent 
and widely spaced cords on the upper shell base without 
intermediate threads, few (usually two) thin threads on the 
subsutural slope, and 16-24 peripheral gemmae on the body 
whorl (usually 22, mean 20.7 at the shell height 18.2-25.9 mm; 
the number of gemmae does not show a strict correlation with 
the shell height). This deep-water species, due to its small size, 
can be confused with young specimens of other species of 
Gemmula. However, the presence of the 'tertiary' apertural 



16 



A.V. SYSOEV 







DEEP-SEA CONOIDEAN GASTROPODS 



17 



notch (characteristic for mature individuals of Gemmula - see 
Kantor & Sysoev, 1991) in one of the shells (H = 20.2) indicates 
that the mentioned shell size characterizes adult specimens of G. 
amabilis. 

Distribution. North-western Indian Ocean, 494-2000 m; also 
probably from north-western Australia, 300^196 m. 

Gemmula (Gemmula) cf. congener (E.A.Smith, 1894) 

Pleurotoma congener E.A.Smith, 1894, p. 160-161, pi. 3,figs4, 5. 
Gemmula congener subspecies congener (E.A.Smith) - Powell, 

1964, p. 251-252, p. 191, figs 1-4; Cernohorsky, 1987, p. 

123-124, figs 1,2-3 (holotype), 4-5. 

Type locality. Bay of Bengal, 128 m. 

Material, stn 176, 1 shell. 

A single broken and heavily worn shell (H = 39.6 mm) can 
probably be referred to G. congener by its very strong 
gemmulated subsutural fold. 

Distribution. Indian Ocean, 198-732 m. 



Subgenus UNEDOGEMMULA MacNeil, 1960 

Type species: Pleurotoma unedo Kiener, 1839—40 (original 
designation) 

Gemmula ( Unedogemmula) unedo (Kiener, 1839-1840) 

Fig. 75 

Pleurotoma unedo Kiener, 1839-1840, p. 19, pi. 14, fig. 1. 
Gemmula (Unedogemmula) unedo (Kiener) - Powell, 1964, p. 

269-270, pi. 175, figs 1, 6, pi. 208, figs 1, 2; Kosuge, 1988, p. 

1 2 1 - 1 22, text-figs 4, 1 3- 1 5, pi. 47, figs 9, 1 0. 
Pleurotoma invicta Melvill, 1910, p. 15, pi. 2, fig. 27. 

Type locality. 'Mers de lTnde' (unedo), Persian Gulf 
{invicta). 

Material, stn 145, 1 specimen. 

The shell from stn 145 is peculiar in the complete absence of 
spiral sculpture on the body whorl. The sculpture is represented 
only by very rough growth lines which is probably a senile 
abnormality. 

Distribution. Persian Gulf to Japan, 73-503 m. 



Subgenus PTYCHOSYRINX Thide, 1925 

I Type species: Pleurotoma (Subulata) bisinuata von Martens, 
1901 (original designation) 



Gemmula ( Ptychosyrinx ) bisinuata (von Martens, 1901) 

Figs 72-74 

Pleurotoma (Subulata) bisinuata von Martens, 1901, p. 17. 
Drillia (Subulata) bisinuata (von Martens) - von Martens, 1903 

[1904], p. 82, pi. 1, fig. 8. 
Ptychosyrinx bisinuata (von Martens) - Thiele, 1925, p. 

176(210), text-fig. 28 (rad.), pi. 46(34), fig. 28 (opera); Powell, 

1964, p. 289-290, pi. 223, figs 1,2. 
Gemmula (Ptychosyrinx) bisinuata (von Martens) 

Cernohorsky, 1987, p. 130, figs 15-17. 

Type locality. 'Valdivia', stn 264, near the coast of Somalia, 
1079 m. 

Material, stn 1 19, 1 specimen and 1 shell; stn 184, 1 specimen. 

The species is rather similar to the closely related G teschi 
(Powell) (see below). The main differences are that, in G 
bisinuata, the subsutural rib is clear, thin, straight or slightly 
wavy, without nodules; the sutures are poorly seen and very 
shallowly impressed. In G teschi, the subsutural rib is very 
weakly developed or absent (or there is a weak to moderate 
subsutural fold), covered with nodules which are the 
continuation of axial folds; the sutures are clear, more or less 
channelled. Additionally, the spiral ribs on the shell base in G. 
bisinuata are clear and prominent; two lower ribs out of three 
upper ones are much stronger than the other. In G teschi, the ribs 
are thinner, more uniform, more numerous and closely spaced; 
the shell base is evenly convex. 

Distribution. East Africa from the Gulf of Aden to 
Malagasy, 818-1463 m. 

Gemmula (Ptychosyrinx) teschi (Powell, 1964) 

Figs 68-70 

Ptychosyrinx timorensis teschi Powell, 1964, p. 291-292, pi. 223, 
figs 5, 6; Abbott & Dance, 1990, p. 238, fig. (holotype). 

Type locality. 'Albatross', stn 5587, NW of Sipadan Id., 
Borneo, 415 fms. 

Material, stn 1 18, 6 specimens. 

Although having been described as a subspecies, G teschi 
obviously warrants specific rank from the fossil G timorensis 
(Tesch, 1915) in having a much broader fusiform shell. The 
specimens from the JME material vary in the prominence of 
spiral ribs on the shell base and the number of peripheral 
tubercles (17-22 on the body whorl, mean 19). In no specimen is 
the subsutural fold as strong and regularly gemmate as in the 
holotype. 

Distribution. Indonesia and Zanzibar, 635-1789 m. 



Figs63-75 Gemmula spp. 63-65 -Gemmula (Gemmula) vagata(E.A. Smith, 1895), stn 1 76 (63, 64) and 188(65), H = 48.9(63), 33.3 (64) and 36.3 
(65) mm; 66, 67, 71 - G (G) amabilis (Jickeli in Weinkauff, 1875), stn 193 (66), 34 (67) and 145 (71), H = 25.8 (66), 23.3 (67) and 21.3 (71) mm; 
68-70 - Gemmula (Ptychosyrinx) teschi (Powell, 1964), stn 1 1 8, H = 28.2 (68), 27.5 (69) and 24.7 (70) mm; 72-74 -G. (P.) bisinuata (von Martens, 
1 901 ), stn 1 1 9 (72) and 1 84 (73, 74), H = 1 8.8 (72) and 30.5 (73, 74) mm; 75 - Gemmula ( Unedogemmula) unedo (Kiener, 1 839^10), stn 145, H = 
95.5 mm. 



18 



A.V. SYSOEV 




DEEP-SEA CONOIDEAN GASTROPODS 
Genus LUCERAPEX hedale, 1936 

Type species: Pleurotoma casearia Hedley & Petterd, 1906 
(original designation) 

Lucerapex adenica Powell, 1964 

Fig. 76 

Lucerapex adenica Powell, 1964, p. 286-287, pi. 221, fig. 3. 

Type locality. 'Mabahiss' (John Murray Expedition), stn 34, 
Gulf of Aden, 1022 m. 

Material, stn 193, 1 shell; also described from stn 34 and 191 
by Powell (1964). 

The specimen figured (H = 31.0 mm which is slightly more than 
in the type specimens) was probably omitted by Powell whose 
original decription of the species was based on the JME 
material. 

Distribution. Gulf of Aden, 274-1080 m. 

Lucerapex molengraaffi (Tesch, 1915) 

Figs 77 & 78 

Pleurotoma (s.str.) molengraaffi Tesch, 1915, p. 28, pi. 77, figs 

54-56. 
Lucerapex molengraaffi (Tesch) - Powell, 1964, p. 287-288, pi. 

220, figs 3, 4, pi. 221, figs 1,2. 

Type locality. Timor, Pliocene. 

Material, stn 145, 1 shell. 

Distribution. Maldive Islands, Borneo, Celebes, Phillipines, 
464-1022 m. 



Family CONIDAE Fleming, 1822 

Subfamily CLATHURELLINAE H. & A.Adams, 1858 

Genus BORSONIA Bellardi, 1839 

Type species: Borsonia prima Bellardi, 1839 (monotypy) 

Borsonia ochracea Thiele, 1925 

Figs 82 & 83 

Borsonia ochracea Thiele, 1925, p. 183-184 (217-218), pi. 38(26), 
figs 1-3, text fig. 26. 

Type locality. 'Valdivia', stn 257, (off Somalia), 1644 m. 

Material, stn 122, 1 specimen; stn 184, 1 shell. 

The JME material agrees well with Thiele's original figures 
differing only in having somewhat narrower and more widely 
spaced spiral ribs (though this may result from rather 
schematized appearance of spiral sculpture in many Thiele's 
figures). The columellar pleat is very weak and can be seen only 



19 

if the aperture is broken. The radular teeth are also very similar 
to those figured by Thiele; the operculum is absent as in the type 
specimens. 

Distribution. East Africa from Zanzibar to the Gulf of Aden, 
693-1 644 m. 



Subgenus CORDIERIA Rouault, 1848 

Type species: Cordieria iberica Rouault, 1850 (subsequent 
designation Cossmann, 1896) 

Borsonia (Cordieria) symbiophora Sysoev, new species 

Figs 5, 12, 13,81 & 85 

Material, stn 26, 4 specimens (paratypes No. 1993103); stn 
118, 6 specimens (holotype No. 1993101 and 5 paratypes No. 
1993102). 

Description of holotype. The shell is of medium size for the 
genus, broadly fusiform, rather stout, strong, white under 
olivaceous periostracum and light-brown inside the aperture, 
and consisting of 7 whorls. The protoconch is missing, and the 
upper whorls are eroded. Definitive whorls are obtusely angled 
at the periphery, slightly concave at the subsutural slope and 
weakly convex below. The body whorl is rather large (0.65 of the 
shell height), the shell base is weakly convex, and passes 
smoothly into the canal. The sutures are wavy and channelled. 
The axial sculpture consists of wide, rounded, short folds which 
are obsolete in the middle part of the subsutural slope (they can 
be traced as very low tubercles just below the suture) and do not 
reach the shell base. The folds are most prominent just below the 
subsutural slope at the whorl periphery. There are 1 2 folds on the 
body whorl and 1 1 on the penultimate. Spiral sculpture is 
represented by wide, rounded, low ribs separated by narrow 
grooves and covering the entire shell surface. On the shell base, 
the ribs become subobsolete, with wider interspaces. The growth 
lines override the ribs making their surface rugose. The aperture 
is rather wide and not differentiated from the wide and short 
canal. The inner lip is smooth, covered by glossy callus. The anal 
sinus is symmetrical, wide and shallow, its deepest part is 
situated in the middle of the subsutural slope. H = 27.4, Hb = 
17.8, Ha = 13.9, D= 11.5mm. 

The paratypes are very variable in the character of spiral 
sculpture which may be either well developed or subobsolete to 
obsolete. There is no correlation between the prominence of 
spiral ribs on the subsutural slope and on the rest part of the 
whorl. Paratypes of smaller size have more biconic shells with 
narrower canals. 

The protoconch is broken in almost all specimens. In the only 
specimen with an intact but eroded protoconch, it seems to 
consist of 1.5 rapidly increasing whorls. 

The radular teeth are typical of borsoniid group of genera, 
small (0.24 mm in paratype from stn 185, H = 21.8 mm), 
straight, rather short, with a cusp at the tooth base. 

The operculum is small, leaf-shaped, vestigial, with a terminal 
nucleus. In one paratype from stn 185 the operculum, probably 



Figs 76-89 Turrinae, Zonulispirinae and Clathurellinae. 76 - Lucerapex adenica Powell, 1964, stn 193, H = 31 .0 mm; 77, 78 - L. molengraaffi 
(Tesch, 1915), stn 145, H = 25.1 mm; 79, 80 - Ptychobela cf. suturalis (Gray, 1838), stn 35 (79) and 188 (80), H = 3 1 . 1 (79) and 22.7 (80) mm; 81, 85 
Borsonia (Cordieria) symbiophora Sysoev, new species, holotype (81) and paratype, stn 1 18, H = 23.8 mm (85); 82, 83 - Borsonia (Borsonia) ochra- 
cea Thiele, 1925, stn 122, H = 37. 0mm;84- Ptychobela cf. nodulosa (Gmdin, 1791), stn 188, H = 29.6mm; 86-88- Typhlomangelia maldivica 
Sysoev, new species, holotype (86) and paratype, stn 143, H = 32.0 mm (87, 88); 89-7: adenica Sysoev, new species, holotype. 



20 



A.V. SYSOEV 



as a result of damage and subsequent repair, is very small, 
subquadrate, with central nucleus (Fig. 12). 

All specimens of B. symbiophora bear actinians on their shells 
and are often entirely covered with them. 

The new species differs from all known Recent species of 
Borsonia, in its stout shell with typically uniform spiral sculpture 
and smooth columella. 

The species is quite comparable with species of Borsonia and, 
especially of the Cordieria subgenus primarily differing in the 
absence of columellar plicae and the presence of operculum. 
However, the prominence (and even presence) of columellar 
plicae can vary greatly among species of the same genus and 
sometimes among shells of the same species. The presence of an 
operculum also cannot be considered as a diagnostic character 
because it is very patchily distributed in the subfamily, where 
repeated and independent reduction and loss of operculum 
undoubtedly occurred, and many faunas demonstrate a full 
range of species with well developed, vestigial or missing 
operculum (e.g. Eastern Pacific - see McLean, 1 97 1 ). 

B. symbiophora is also similar to species of the subgenus 
Borsonellopsis McLean, 1971 of the genus Borsonella Dall, 1890. 
The type species of Borsonellopsis, Leucosyrinx erosina Dall, 
1908, possesses similar sculpture and shell outlines as well as a 
vestigial operculum, and lacks columellar plicae. On the other 
hand, it differs considerably from Borsonella s.str. and may not 
be congeneric. 

Distribution. Gulf of Aden and off Mombasa (Kenya), 
1789-23 12 m. 



Genus BA THYTOMA Harris & Burrows, 1 89 1 

Type species: Murex cataphractus Brocchi, 1814 (monotypy) 

Subgenus PARABATHYTOMA Shuto, 1961 

Type species: Pleurotoma striatotuberculata Yokoyama, 1928 
(original designation) 

The differences between the generally accepted subgenera of 
Bathytoma (see Powell, 1966) seem to be rather slight. Kilburn 
( 1 986) mentioned that Parabathytoma differs from Micantapex 
in having radular teeth without an elongate base and in the 
absence of brephic arcuate riblets at the place of the protoconch 
transition into teleoconch whorls. However, in all the species 
described below, the presence of long, curved teeth without an 
elongate base is associated with the presence of arcuate riblets at 
the place of the protoconch termination, i.e. these species 
possess characters of both Parabathytoma and Micantapex 
sensu Kilburn. Thus, the only feature distinguishing these 
subgenera is the shape of radular teeth. The question is further 
complicated by the fact that the type species of Parabathytoma is 
a fossil Pleurotoma striatotuberculata Yokoyama, 1928, while the 
radula of type species of Micantapex, Bathytoma agnata Hedley 
& Petterd, 1906, is unknown. Nevertheless, the species listed 
below are provisionally included into Parabathytoma on the 
basis of their radular morphology. 



Bathytoma (Parabathytoma) prodicia Kilburn, 1986 

Figs 90 & 91 

Bathytoma (Parabathytoma) regnans (non Melvill, 1918) - 

Kilburn, 1971, p. 31, figs 2c, 2f, 4b 
Bathytoma (Parabathytoma) prodicia Kilburn, 1986, p. 643, figs 

22-23. 



Type locality. East of 
Mozambique), 300-310 fms. 

Material, stn 119, 1 shell. 



Bazaruto Island (Southern 



The shell from the JME material differs from the holotype 
figured by Kilburn in having less prominent peripheral nodules, 
shallower anal sinus, and flattened shell base. However, all other 
essential conchological characters including the shell 
proportions (D/H = 0.43, Ha/H = 0.52) are similar to B. prodicia. 
An additional smaller (H = 23.5 mm) specimen collected off 
Zanzibar by R/V 'Vityaz' (stn 4680, 740 m) is in some respects 
intermediate between the typical B. prodicia and the JME shell. 

Distribution. Southern Mozambique to Zanzibar, 420-1463 
m. 

Bathytoma (Parabathytoma) oldhami (E.A.Smith, 1899) 

Figs 7 & 92-93 

Pleurotoma (Bathytoma) oldhami E.A.Smith, 1899, p. 238. 
Pleurotoma oldhami E.A.Smith -Alcock&McArdle, 1901, pi. 9, 
figs 2, 2a. 

Type locality. 'Investigator', stn 229, off Travancore coast, 
360 fms. 

Material, stn 145, 1 specimen. 

The JME specimen is of approximately the same size as the 
holotype and very similar to the figure of the latter, differing 
only in slightly broader shell (H = 41.0 mm, D = 15.8 mm vs. 43 
and 15 mm in the holotype). The protoconch consists of about 
1.5 smooth globose whorls followed by several arcuate axial 
riblets which gradually become stronger and pass into the 
teleoconch sculpture. The radular teeth (Fig. 7) are long and 
strongly curved, of typical shape for the subgenus. The mean 
tooth lengtrris 0.77 mm. 

Distribution. Southern India and Maldive Islands, 494-658 
m. 



Bathytoma (Parabathytoma) regnans Melvill, 1918 

Figs 8, 16 & 94-97 

Bathytoma regnans Melvill, 1918, p. 68, textfig. 
Bathytoma regnans Melvill - Kilburn, 1986, p. 718, fig. 168 
(holotype). 

Type locality. Indian Ocean, Investigator Expedition 
(probably the Bay of Bengal). 

Material, stn 34, 1 specimen and 5 shells; stn 188, 2 shells; stn 
193, 6 shells. 






Figs 90-101 Clathurellinae. 90, 91 - Bathytoma (Parabathytoma) prodicia Kilburn, 1986, stn 1 19, H = 32.3 mm; 92, 93 - B. (P.) oldhami^. .A. 
Smith, 1 899), stn 145, H = 41 .0 mm; 94-97 - B (P.) regnans Melvill, 1918, stn 188 (94, 97), 34 (95) and 193 (96), H = 27.5 (94), 26.6 (95), 24.4 (96) 
and 25.5 (97) mm; 98-100- B. (P)fissa (von Martens, 1901), stn 176, H = 35.3 (98), 34.7 (99) and 38.4 (100) mm; 101 - Typhlosyrinx praecipua 
(E.A. Smith, 1 899), stn 1 84, H = 30.2 mm. 



DEEP-SEA CONOIDEAN GASTROPODS 



21 




22 



A.V. SYSOEV 



The species is rather variable in the shell proportions: the shell 
may be elongated (Fig. 94) and very similar to the holotype 
figured by Kilburn (1986), or more stout and broad (Figs 95 & 
96). The H/D ratio varies from 2.03 to 2.55 (2.44 in the 
holotype). The prominence of the columellar pleat can also vary 
up to almost complete absence. However, there are rather 
constant features which are characteristic of the species and 
distinguish it from B. fissa (see below). These are a strongly 
projecting peripheral keel, distant primary spiral cords with a 
thin secondary rib in most interspaces, and low position of the 
peripheral keel, which is bordered or almost bordered by a 
deeply channelled suture even on the last spire whorls. The 
protoconch is similar to that of B. oldhami. Radular teeth are 
typical for the subgenus, awl-shaped, curved, and rather short. 
The mean tooth length is 0.46 mm at H = 26.6 mm. The 
operculum is small, oval, with subterminal nucleus. 

Distribution. Gulf of Aden and, probably, the Bay of Bengal, 
528-1080 m. 

Bathytoma ( Parabathytoma) fissa (von Martens, 1901) 

Figs 9, 17 & 98-100 

Pleurotoma ( Dolichotoma) fissa von Martens, 1901, p. 18. 
Genota (Dolichotoma) fissa -von Martens, 1903 [1904], p. 87, pi. 
1, fig. 14. 

Type locality. 'Valdivia', stn 264 (Somalia), 1079 m. 

Material, stn 176, 2 specimens and 12 shells. 

The species differs from B. regnans in having closely set spiral 
cords, with almost equally strong granular secondary cords in 
the interspaces, and less prominent peripheral keel which is 
situated rather high on last spire whorls. The sutures are less 
distinctly channelled than in B. regnans. The shell proportions 
and the prominence of peripheral keel may vary (Figs 98-100). 
There are one to three more or less developed columellar pleats; 
usually two pleats are present and the upper one may be 
subdivided into two by a groove. The radula and operculum are 
similar to those of B. regnans. The mean tooth length is 0.51 mm 
at H = 35.3 mm. 

Shells from an additional sample (R/V 'Akademik Mstislav 
Keldysh', stn 1089, Tajoura Rift, Gulf of Aden, 857-900 m) 
show the same characteristic features and the same range of 
variability but are of larger size (up to 42.5 mm in height) and 
possess 1 (usually) or 2 columellar pleats. 

Distribution. Somalia and Gulf of Aden, 665-1079 m. 



Genus TYPHLOMANGELIA G.O.Sars, 1878 

Type species: Pleurotoma nivale Loven, 1846 (monotypy) 

Typhlomangelia adenica Sysoev, new species 

Figs 4, 14&89 

Material, stn 26, 1 shell (paratype No. 1993106); stn 185, 2 
specimens (holotype No. 1993104 and paratype No. 1993105). 

Description of holotype. The shell is small, broad, turreted, 
solid, covered by thin light-brown periostracum, and consists of 
4 remaining whorls. Protoconch and probably some upper 
whorls are missing. The whorls are strongly angled at the 
periphery, the subsutural slope is concave. There is a weak 



subsutural fold. Sutures are very shallow and indistinct. The 
body whorl is large, and the shell base is weakly convex and not 
differentiated from the canal. The axial sculpture consists of 
numerous narrow oblique folds reaching the lower suture and 
abruptly disappearing on the subsutural slope. The folds form 
pointed tubercles just below the subsutural slope. On the body 
whorl, they rapidly weaken downwards and do not reach the 
shell base. There are 19 folds on the body whorl and 15 on the 
penultimate. The subsutural slope is smooth except for growth 
lines. A single spiral cord is situated on the subsutural fold, 2-4 
cords are present below the subsutural slope on the spire whorls, 
and there are about 1 5 low, widely and evenly spaced cords on 
the body whorl, which become weaker towards the anterior end 
and finally disappear. The aperture is oval, not differentiated 
from the straight and short canal. The inner lip is covered with a 
white callus which is thickened and becomes distinctly bordered 
on the canal. The anal sinus is deep, broad and rounded, slightly 
asymmetrical, its deepest point is situated just below the middle 
of subsutural slope. H = 10.2, Hb = 6.6, Ha = 4.7, D = 5.6 mm. 

The paratypes are very similar to the holotype but smaller (H 
= 7.7, D = 4.5 in paratype from stn 185 and H = 6.9, D = 4.1 in 
paratype from stn 26). The operculum is rather large in 
comparison to the aperture size, broadly leaf-shaped, with a 
terminal nucleus. Radular teeth are typical of the genus, 
awl-shaped, with short and straight shaft and relatively broad 
base, 0.2 mm in length. 

The species differs from all other species of the genus in its 
very small, broad shell, with a very short and straight canal. 

Distribution. Gulf of Aden, 2000-2312 m. 

Typhlomangelia maldivica Sysoev, new species 

Figs 6, 15 & 86-88 

Material, stn 143, 7 specimens (holotype No. 1993107 and 6 
paratypes No. 1993108). 

Description of holotype. The shell is elongate fusiform, 
slender, with a high spire, rather solid, covered with thin 
grayish-brown periostracum, and consists of protoconch and 9 
teleoconch whorls. The protoconch consists of about two 
globose smooth whorls, the surface is partly eroded. Definitive 
whorls are angled at the periphery, concave on the subsutural 
slope, with distinct subsutural fold. The body whorl occupies 
0.58 of the shell height, the shell base is weakly convex and 
passes smoothly into the slightly twisted canal. The sutures are 
distinctly channelled, especially in last whorls. The axial 
sculpture consists of rounded folds below the subsutural slope; 
they are most prominent and tuberculate in their uppermost 
parts and weaken towards the lower suture. The folds tend to 
become smoother on last spire whorls, and on the body whorl 
they are very low and restricted to the whorl periphery. There are 
13 folds on the body whorl and 12 on the penultimate. Spiral 
sculpture is represented by rather strong and widely spaced 
cords. One (or two on last whorls) strong cord is situated on the 
subsutural slope, 1-3 cords are developed below the subsutural 
slope of spire whorls ( 1 on initial whorls and 3 on the last one), 
and about 20 cords cover the body whorl and the canal. On the 
body whorl periphery, a weaker cord is situated in each 
interspace between primary cords. The subsutural slope of the 
initial whorls is smooth, but on subsequent whorls 1-5 thin 
riblets are developed, with a stronger one in the centre of the 
slope. The aperture is rather narrow, oval. The canal is 
moderately long. The anal sinus is deep, rounded, slightly 



DEEP-SEA CONOIDEAN GASTROPODS 



23 



asymmetrical, with the apex situated just below the middle of the 
subsutural slope. H = 30.2, Hb = 17.4, Ha = 13.9, D = 8.8 mm. 

The paratypes vary in details of spiral sculpture, especially on 
the subsutural slope, where several rather strong and evenly 
spaced riblets may be developed. In some paratypes the spiral 
cords are more numerous, closely spaced on the canal and more 
or less irregularly distributed over the body whorl surface. There 
may be from 13 to 15 axial folds on the body whorl. The largest 
paratype has H = 32.0 and D = 9.1 mm. 

Operculum is small, oval, with terminal nucleus. Radular 
teeth are long and narrow, more or less curved, without a solid 
base, rather large (0.74 mm on the average at H = 27.8 mm). 

The new species resembles the type species of Typhlomangelia, 
Pleurotoma nivale Loven, 1846, differing well in the elongate 
shell with a high spire and prominent spiral ribs. 

Distribution. Maldive Islands, 797 m. 



Genus TYPHLOSYRINX Thiele, 1925 

Type species: Pleurotoma (Leucosyrinx) vepallida von Martens, 
1902 (original designation) 

Typhlosyrinx praecipua (E.A.Smith, 1899) 

Fig. 101 

Pleurotoma (Surcula) praecipua E.A.Smith, 1899, p. 239; 

Annandale & Stewart, 1 9 1 0, pi. 2 1 , figs 4, 4a. 
Typhlosyrinx praecipua (E.A.Smith) - Powell, 1969, p. 360-361 , 

pi. 272, figs 2, 3. 

Type locality. 'Investigator', stn 229, off Travancore coast 
(India), 360 fms. 

Material, stn 184, 1 shell. 

The shell from stn 184 (H = 30.2 mm) corresponds quite well 
with the description and illustration of the type specimen. Axial 
folds (1 1 on the penultimate whorl) abruptly disappear on the 
border between the penultimate and body whorls. The spiral 
sculpture is developed only on the whorl periphery and the shell 
base, and consists of low, broad, flattened, wavy riblets unequal 
in width and separated by narrow grooves. 

Distribution. India and Gulf of Aden, 658-1270 m. 



Genus GLYPHOSTOMA Gabb, 1872 

Type species: Glyphostoma dentiferum Gabb, 1872 (monotypy) 

Glyphostoma maldivica Sysoev, new species 

Figs 1 02 & 103 

Material, stn 145, 1 shell (holotype No. 1993109). 

Description. The shell is rather small, thick, solid, 
yellowish-white, and consists of the protoconch and 5 3/4 
teleoconch whorls. The protoconch consists of 3 whorls; its 
initial part is represented by 1.5 rapidly increasing 
semi-transparent, smooth whorls followed by more solid angled 
whorls sculptured below the periphery by a narrow keel. The 
teleoconch whorls are concave below the suture and angled at 
the periphery. The sutures are clear, shallow, and wavy. The shell 
base is weakly convex and passes smoothly into the canal. The 



shell surface is distinctly and minutely granular, the granulation 
is better seen in the interspaces between the axial folds. The 
growth lines are mostly indistinct. The axial sculpture is 
represented by rounded folds extending from the subsutural 
slope to lower suture and, on the body whorl, to the canal. There 
are 24 such folds on the body whorl and 19 on the penultimate. 
On the subsutural slope, the axial sculpture consists of 
numerous curved and rather weak folds with sharpened crests. 
They generally represent the continuation of main axial, folds 
but there may also be interstitial folds; as a result, the subsutural 
slope of the body whorl is covered with 32 folds. The spiral 
sculpture is represented by strong ribs almost equal in 
prominence to the axial folds. At the intersection with axial 
sculpture, the ribs form rounded tubercles. The interspaces 
between spiral ribs are covered by closely set threads. The ribs on 
the subsutural slope are much smaller corresponding to much 
smaller axial folds. The aperture is elongate-oval. The inner lip 
bears two rather prominent pointed tubercles in its middle part 
and several smaller ones in the lower part. The outer lip is 
sharpely and wavely edged and bordered by a heavy curved 
varix. The inner surface of the aperture bears one strong 
tubercle in the upper part and a group of 5 tubercles below. The 
anal sinus is deep, U-shaped, bordered with callus and 
constricted at its entrance by a heavy tuberculated callus pad. 
The canal is straight along most its length and slightly curved 
backwards near the end. H = 17.6, Hb= 11.8, Ha = 9.8, D = 8.6 
mm. 

In general outline the new species is most similar to 
Clathurella perlissa E.A.Smith, 1904 from the Andaman Islands 
but differs in the character of sculpture and apertural 
armament. The species corresponds well to the genus 
Glyphostoma in all important conchological characters such as 
characteristic protoconch, granular surface, prominent 
subsutural slope with different sculpture (in contrast to Etrema), 
and strong intersecting spiral and axial ribs. 

Distribution. Maldive Islands, 494 m. 

Glyphostoma supraplicata Sysoev, new species 

Figs 1 04 & 105 

Material, stn 176, 6 shells (holotype No. 1993110 and 5 
paratypes No. 1993111). 

Description of holotype. The shell is rather small, slender, 
fusiform, relatively solid, white, consists of protoconch and 
almost 7 teleoconch whorls. The protoconch consists of 3.5 
whorls, the tip is small and papillate. Initial embryonal whorls 
are smooth, the two last whorls angled, with a cord-like 
peripheral keel, the second weaker keel encircles the lower 
suture. The spire is tall, occupies about 0.4 of the shell height. 
Definitive whorls are angled at the shoulder and in the lower 
part, the subsutural slope is weakly concave or almost flat. The 
sutures are clear, shallow, slightly wavy. The entire shell surface is 
densely and minutely granulated, but the granulation more or 
less disappears on prominent parts of the sculpture. Growth 
lines are thin and irregularly elevated. Axial sculpture consists of 
strong, oblique, rounded folds separated by interspaces 
approximately equal in width to the folds. The folds reach the 
canal, gradually weakening on the shell base. On the subsutural 
slope, they are much weaker, narrower, almost obsolete near the 
upper suture, and curved in correspondence to the anal sinus 
scars. There are 15 folds on the body whorl and 13 on the 
penultimate. Spiral sculpture is represented by broad cords 



24 



A.V. SYSOEV 




DEEP-SEA CONOIDEAN GASTROPODS 



25 



below the shoulder (2 on the spire whorls and 18 on the body 
whorl plus canal). The cords are separated by 2-3 times wider 
intervals. They are often subobsolete in interstices between axial 
folds but very strong when overrided the latter. On the body 
whorl, the cords gradually diminish in width and prominence 
towards the canal end. There are \-4 (usually 2) thin threads in 
the interspaces between cords, except between those on the 
canal, and 3-4 low rounded threads on the subsutural slope. 
Strong rounded tubercles are formed at the intersection between 
axial folds and spiral cords resulting in a beaded appearance of 
the spiral sculpture. The aperture is rather narrow, gradually 
narrowing towards the canal. The inner lip bears about 13 weak 
transverse plicae in its lower part, the plicae become closer to 
each other towards the canal end. There is a moderately 
developed parietal callus pad which is weakly tuberculate in its 
lower part. The outer lip has a sharpened edge, with a strong 
varix behind. Inside the aperture, there are 5 transverse plicae 
most prominent in the region corresponding to the varix, and a 
strong tubercle just below the anal sinus. The sinus is 
semi-tubular, U-shaped, broad and rounded, directed outside in 
relation to subsutural slope. The canal is long, slightly twisted, 
and obliquely truncated at the end. H = 22.6, Hb = 13.4, Ha = 
11.3, D = 7.9 mm. 

Paratypes vary in minor details of sculpture, e.g. in the 
prominence of axials on the subsutural slope, and in more or less 
strong nodules at the intersections between spiral and axial 
elements. The largest paratype is 22.6 mm high (protoconch 
missing). 

The new species is quite similar to G sultana (Thiele, 1925) 
from East Africa, differing in the more slender shell (H/D ratio is 
2.70-2.91 vs. 2.55 in the holotype of G. sultana) and in the 
presence of weakened but distinct axial folds on the subsutural 
slope. A peculiar feature of the new species is the presence of a 
second keel encircling the suture on last protoconch whorls. 

Distribution. Gulf of Aden, 655-732 m. 



Subfamily DAPHNELLINAE Deshayes, 1863 
Genus CRYPTODAPHNE Powell, 1942 

Type species: Cryptodaphne pseudodrillia Powell, 1942 (original 
designation) 

Cryptodaphne gradata (Schepman, 1913) 

Fig. 108 

Pleurotomella gradata Schepman, 1913, p. 445, pi. 30, fig. 2. 
Cryptodaphne gradata (Schepman) -Shuto, 1971, p. 1 1, pi. 2, figs 
4-6. 

Type locality. 'Siboga', stn 159, Halmahera Sea, 411m. 

Material, stn 176, 1 shell. 

The shell from the JME material basically conforms to the 
description and illustrations of the holotype given by Schepman 
and Shuto (see synonymy). However it differs from the latter in 
some remarkable characters. The shell is much larger than that in 



the holotype (18.5 mm vs. 10.0 mm), with a proportionally 
higher spire, at approximately the same number of teleoconch 
volutions (6+ vs. 6). The anal sinus is deeper, with its apex 
situated lower on the subsutural slope. The initial three whorls 
of the teleoconch bear oblique axial folds below the peripheral 
angulation. These folds form nodes on the angulation, making it 
crenulated, and rapidly weaken towards the lower suture. On 
subsequent whorls, they become less developed and disappear 
on the body whorl. The spiral sculpture lacks a regular 
alternation of strong and weak spirals mentioned by Shuto. The 
sculpture has a cancellated appearance due to the intersection 
with growth lines. Also there are widely spaced, thin, and weak 
but distinct spiral threads on the subsutural slope which were 
not described by either Schepman or Shuto. 

Nevertheless, these differences are not essential and seemingly 
do not extend beyond the range of intraspecific and 
geographical variability of C. gradata. 

Distribution. Halmahera Sea and Gulf of Aden, 41 1-732 m. 



Genus FAMELICA Bouchet & Waren, 1980 

Type species: Pleurotomella catharinae Verrill & Smith, 1884 
(original designation) 

Famelica tajourensis Sysoev & Kantor, 1987 

Fig. 112 

Famelica tajourensis Sysoev & Kantor, 1987, p. 1257, fig. g, d, e, 
zh. 

TYPE LOCALITY. 'Akademik Kurchatov', stn 1095, Tajoura 
Rift, Gulf of Aden, 1330-1406 m. 

Material, stn 188, 1 shell. 

The JME shell agrees well with the type material and differs 
mainly in having thin and transparent shell walls compared with 
rather thick and solid in the type specimens. 

The species is rather different from the type species of the 
genus Famelica, but very similar to F. monotropis (Dautzenberg 
& Fischer, 1896), which was included into Famelica by the 
authors of the genus (Bouchet & Waren, 1980). On the other 
hand, the genus seems to be rather heterogeneous in respect to 
species originally included into it. Nevertheless, I do not know 
any other genus which can accomodate F. tadjourensis. A similar 
genus is Pagodidaphne Shuto, 1983, but it differs in having a less 
distinct keel on the whorl shoulder and much less elongate body 
whorl with a short canal. 

Distribution. Gulf of Aden, 528-1406 m. 



Genus GYMNOBELA Verrill, 1884 

Type species: Gymnobela engonia Verrill, 1884 (subsequent 
designation Cossmann, 1896) 



Figs 102-1 12 Clathurellinae and Daphnellinae. 102, 103 - Glyphostoma maldivica Sysoev, new species, holotype; 104, 105 - G. supraplicata Sysoev, 
new species, holotype; 106, 107 - Gymnobela adenica Sysoev, new species, holotype (106) and paratype, stn 1 85, H = 7. 1 mm (107); 108 - Crypto- 
daphne gradata (Schepman, 1913), stn 178, H = 18.5 mm; 109-111 - Gymnobela africana Sysoev, new species, holotype (109, 110) and paratype, stn 
1 1 8, H = 68.0 mm (1 1 1); 1 12 - Famelica tajourensis Sysoev & Kantor, 1987, stn 1 88, H = 10.6 mm. 



26 



A.V. SYSOEV 



Gymnobela adenica Sysoev, new species 

Figs 1 06 & 107 

Material, stn 185, 2 specimens (holotype No. 1993112 and 
paratypeNo. 1993113). 

Description of holotype. The shell is small, broadly biconic, 
thin, yellowish-white, and consisting of 5 remaining whorls. The 
protoconch is missing, and the upper teleoconch whorls are 
eroded. The whorls are angled below the periphery. The 
subsutural slope is almost flat on the upper spire whorls and 
concave on the body whorl. The uppermost part of subsutural 
slope is slightly raised forming an indistinct subsutural fold. The 
sutures are shallow. Growth lines are mostly indistinct, some of 
them form clear, narrow, oblique folds regularly set on the upper 
third of the subsutural fold and approximately twice as 
numerous as main axial folds. The latter are strongly oblique, 
narrow, with sharpened crests, and tuberculate at the place of 
whorl angulation. The folds are separated by narrow intervals, 
abruptly disappear on the subsutural slope and extend to the 
lower suture on the spire whorls and to the upper part of the 
shell base. There are 21 folds on the body whorl and 20 on the 
penultimate. Spiral ribs (about 30 on the body whorl plus canal) 
are strong, flattened and uniform except for narrower ones on 
the canal. Intervals between the ribs are approximately equal to 
ribs in width. The subsutural slope is densely covered with thin, 
low, rounded and closely set riblets (about 14 on each of two last 
whorls). The shell base with a distinct bend passes into a short 
and straight canal. The aperture is rather small, subrectangular, 
with the inner lip distinctly bent. The canal is narrow, attenuated 
at its end. The anal sinus is moderately deep, broadly rounded, 
its deepest point is situated in the middle of subsutural slope. H 
= 9.5, Hb = 6.9, Ha = 5.1, D = 6.0 mm. 

The paratype is very similar to the holotype except for smaller 
size (H = 7.1, D = 5.3 mm, 3 teleoconch whorls), smaller H/D 
ratio, and longer folds formed by growth lines which often 
occupy the whole subsutural slope. There are 24 main axial folds 
on the body whorl. The last protoconch whorl preserved is 
covered with typical diagonally cancellated sculpture. 

The species differs from other species of the genus in its small 
broadly biconic shell with numerous strongly oblique axials and 
short attenuated canal. 

Distribution. Gulf of Aden, 2000 m. 



Subgenus BATHYBELA Kobelt, 1905 

Type species: Thesbia nudator Locard, 1897 (subsequent 
designation Dall, 1918) 

Gymnobela ( Bathybela) africana Sysoev, new species 

Figs 109-1 11 

Material, stn 118, 1 specimen (holotype, No. 1993114) and 1 
shell (No. 1993115). 

Description of holotype. The shell is large, broadly 
fusiform, thin but solid, reddish-brown to light-brown, and with 
a slightly glossy surface. It consists of 6 whorls, the protoconch is 
missing. The whorls are obtusely angled above the periphery, 
moderately convex below the angulation and slightly concave 
above it. The sutures are rather shallowly impressed, clear, 
straight or wavy in some places. The body whorl is large, 



occupies about 0.7 of the shell height; the shell base is weakly 
convex, with a distinct bend passes into the straight canal. The 
axial sculpture consists of narrow oblique folds, often with 
sharpened crests. The folds begin in the lower part of the 
subsutural slope and extend to the lower suture on the spire 
whorls and only to the periphery on the body whorl. They are 
most prominent in the place of the whorl angulation. There are 
1 7 folds on the body whorl and 1 5 on the penultimate one. The 
growth lines are indistinct on the whorl surface except the 
subsutural slope, some of them are thickened and raised. The 
thickened growth lines, however, do not form the regularly 
arranged plicae on the subsutural slope which are characteristic 
of many deep-sea Daphnellinae. The spiral sculpture is 
represented by rather wide, flattened, wavy ribs unequal in size 
and separated by narrow grooves. The subsutural slope is 
smooth except for several very feeble spiral lines. The aperture is 
broad, the inner lip forms a distinct bend, and the columella is 
twisted. The anal sinus, judging from the growth lines, is shallow 
and broad, its deepest point is situated immediately above the 
middle of subsutural slope. The dried soft body has no 
operculum. 

H = 54.7, Hb = 38.5, Ha = 30.1, D = 22.7 mm. 

The shell of the paratype is larger than in the holotype (H = 
68.0, Hb = 48.2, Ha = 38.5, D = 30.0 mm). It was dead-collected 
and the surface is rather worn. The paratype differs from the 
holotype mainly in its shorter axial folds developed on the body 
whorl only at the place of angulation and obsolete in the last 
quarter of body whorl, and in the canal curved backwards. 

The new species is rather similar to Spergo sibogae Schepman, 
1913 from Indonesia differing in much broader shell (H/D = 3.0 
in S. sibogae and 2.3-2.4 in G. africana). It also shows some 
similarity to Pontiothauma pacei E.A.Smith, 1906 from India 
and Ceylon differing in somewhat more slender shell with short 
axial folds and faint spiral sculpture. 

Distribution. East Africa eastward of Mombasa, 1 789 m. 



Subgenus THETA Clarke, 1959 

Type species: Pleurotomella (Theta) lyronuclea Clarke, 1959 
(original designation) 

Gymnobela (Theta) daphne lloides (Dall, 1895) 

Figs 1 13 & 114 

Mangelia (Spergo) daphnelloides Dall, 1895, p. 683-684, pi. 31, 
fig. 11. 

Type locality. 'Albatross', stn 3476 (Hawaiian Islands), 298 
fms. 

Material, stn 118, 1 shell. 

The species was originally described within the subgenus Spergo 
which is considered as a full genus by all modern authors. 
However the species differs from the type species of Spergo, 
Mangilia (Spergo) glandiniformis (Dall, 1895), in all important 
conchological characters included by Dall in the original 
diagnosis of the subgenus (the latter was based to a considerable 
extent on the soft body characteristics which are presently 
treated as insignificant for taxonomy at the generic level), i.e. in 
having well developed sculpture and rather deep anal sinus. On 
the other hand, the species is very similar to the group of species 
assigned by Bouchet & Waren (1980) to the genus Theta. The 



DEEP-SEA CONOIDEAN GASTROPODS 



27 




Figs 113-122 Daphnellinae and Mangeliinae. 113, 114 -Gymnobela (Theta) daphnelloides (Dall, 1895), stn 118, H = 27.0; 115- Mioawateria exten- 
saeformis (Schepman, 1913), stn 26, H = 9.9 mm; 116, 117- Xanthodaphne maldivica Sysoev, new species, holotype; 118-121 - Benthomangelia 
brachytona (Watson, 1881), stn 1 19, H = 17.1 mm (118, 119) and lectotype, BM(NH) 1887.2.9.1034, H = 15.1 mm (120, 121); 122- B. trophonoidea 
Thiele, 1925, stn 185, H = 10.0 mm. 



latter, in its turn, is too poorly distinguished from Gymnobela to 
warrant a generic status and should be regarded as a subgenus. 

Distribution. Hawaiian Islands and East Africa, 545-1789 m. 
The present record is a great range extension for the species. 



Genus MIOAWATERIA Vella, 1954 

Type species: Awateria personata Powell, 1954 (original 
designation) 



Mioawateria extensaeformis (Schepman, 1913) 

Fig. 115 

Pleurotomella extensaeformis Schepman, 1913, p. 446, pi. 30, fig. 

5. 
Gymnobela extensaeformis (Schepman) - Thiele, 1925, p. 

490(224), pi. 41(29), fig. 9. 
Magnella extensaeformis (Schepman) - Shuto, 1971, p. 15-16, 

pi. 2, figs 10-12. 

Type locality. 'Siboga', stn 212, Banda Sea, 462 m. 



28 



A.V. SYSOEV 



Material, stn 26, 2 specimens. 

Of the two specimens available (H = 9.9 and 6.3 mm, both 
without protoconch), the smaller is comparable in size to those 
illustrated by Schepman (8.5 mm, or 8.0 mm according to Shuto, 
1971) and Thiele (6.25 mm) and quite similar to them. The larger 
specimen (Fig. 115) differs in having a broader canal, less 
excavated subsutural slope, and much weaker peripheral spiral 
keel on the body whorl. The spiral sculpture of both specimens is 
subobsolete on the body whorl periphery and upper part of the 
shell base becoming stronger towards the canal. This is in 
contrast to both Schepman's and Thiele's figures, but agrees well 
with Shuto's (1971) illustration of the holotype. 

The same species was probably figured by Thiele (1925) as the 
North- Atlantic Gymnobela extensa (Dall, 1881). Thiele's figure, 
apparently based on a specimen from Sumatra, differs from his 
illustration of M. extensaeformis in having obsolete spiral ribs 
on the body whorl except for those on the lower shell base and 
canal, but, as mentioned above, this is the feature characteristic 
of M. extensaeformis. 

Recently this species was assigned by Shuto (1971) to 
Magnella Dittmar, 1960. Later, Maxwell (1988) synonymized 
the latter genus with the New Zealand genus Mioawateria Vella, 
1954. This synonymization seems to be reasonable, though the 
status of Mioawateria itself remains somewhat uncertain due to 
great similarity to Gymnobela Verrill, 1884. The main difference 
between two latter genera is the shape of anal sinus which is very 
shallow in Mioawateria. At the same time, the shape of sinus in 
Gymnobela is rather variable, and there are species, traditionally 
included into Gymnobela, quite comparable to Mioawateria in 
this character (e.g. G. blakeana (Dall, 1881)). Thus, after 
comparative examination of broad range of Gymnobela species 
in respect to the anal sinus shape, Mioawateria may be either a 
large and very widely distributed genus or a synonym of 
Gymnobela. 

Distribution. East Africa (Gulf of Aden to Kenya), Sumatra, 
Banda Sea, 439-2312 m. The present record is the deepest one. 



Genus XANTHODAPHNE Powell, 1942 

Type species: Pleurotoma (Thesbia) membranacea Watson, 1886 
(original designation) 

Xanthodaphne maldivica Sysoev, new species 

Figs 1 16 & 117 

Material, stn 143, 1 specimen (holotype No. 1993116). 

Description. The shell is narrowly fusiform, slender, thin but 
solid, light-brown, with glossy surface, consists of the 
protoconch and 9 teleoconch whorls. The protoconch is partly 
broken off, but the remaining 1.5 whorls are covered with the 
typical diagonally cancellated sculpture. The teleoconch whorls 
are slightly concave in the upper part and weakly convex below. 
Early teleoconch whorls are angled in the lower part just near the 
suture, this angulation rapidly shifts upwards and becomes less 
prominent and practically disappears on the 6th whorl. There is 
a weak subsutural fold on initial teleoconch whorls. The sutures 
are very shallow, distinct, and more or less straight. The growth 
lines are clear, thin, numerous, and strongly curved. The 
sculpture is represented only by low, wide ribs on the canal; these 
rapidly become obsolete on the shell base. The shell base is 
weakly convex and smoothly passes into the canal. The canal is 



straight, not differentiated from narrow aperture. The inner lip is 
covered with a very weak callus which becomes thicker towards 
the canal extremity. The anal sinus is wide and moderately deep, 
subsutural, 'reversed L'-shaped, its deepest part is situated just 
below the suture. The outer lip strongly projects forward below 
the sinus. H = 29.0, Hb = 17. 1, Ha = 14.3, D = 9.0 mm. 

The new species differs from all known species of the genus in 
its slender and narrow shell almost completely devoid of spiral 
sculpture. 

Distribution. Maldive Islands, 797 m. 



Subfamily MANGELIINAE Fischer, 1883 
Genus BENTHOMANGELIA Thiele, 1925 

Type species: Surcula trophonoidea Schepman, 1913 (original 
designation) 

Benthomangelia brachytona (Watson, 1881) 

Figs 118-121 



Pleurotoma (Drillia) brachytona Watson, 1881, p. 415. 
Pleurotoma (Spirotropis) brachytona Watson - Watson, li 
324-325, pi. 18, fig. 3. 



\ p. 



Type locality. 'Challenger', stn 191, off the Arrou Island, 
south-west of Papua, 800 fms. 

Material, stn 119, 1 shell. 

The specimen from the stn 1 19 (H = 17.1 mm) is quite similar to 
Watson's illustration and to the photograph of one of two 
syntypes (which should be designated as lectotype) (Figs 120 & 
121, H = 15.3 mm according to Watson) (the second syntype is 
represented by broken and heavily worn shell) stored in the 
NHM. It differs from the lectotype only in more slender shell 
(H/D = 2.20 vs. 2.03 (Watson's measurements) or 1 .96 (measured 
by the photograph) in Watson's specimen) with fewer axial folds. 

Distribution. East Africa and Indonesia, 1463 and 
1207-1463 m. 

Benthomangelia trophonoidea (Schepman, 1913) 

Fig. 122 

Surcula trophonoidea Schepman, 1913, p. 62(426)-63(427), pi. 

28, fig. 3. 
Mangelia (Benthomangelia) trophonoidea (Schepman) - Thiele, 

1925, p. 190(224)-191(225), pi. 27(39), fig. 25, text-fig. 25. 
Benthomangelia trophonoidea (Schepman) - Okutani, 1966. p. 

23, text-fig. 11. 
? Marshallena gracilispira Powell, 1969, p. 370-371, pi. 281, fig. 2. 

Type locality. 'Siboga', stn 45, Flores Sea, 794 m. 

Material, stn 185, 1 shell. 

The shell from the JME material is small (H = 10.0 mm, which is 
smaller than all the previously recorded specimens, i.e. 16 mm 
(Schepman), 1 5.9 mm (Thiele, measured by the figure), and 15.5 
mm (Okutani, measured by the figure)) and apparently rather 
young. It differs from the original description and figure in 
having prominent tubercles on the subsutural fold of all the shell 
whorls and in less curved and more developed axial ribs on the 
subsutural slope. The presence of short plicae on early whorls, 



DEEP-SEA CONOIDEAN GASTROPODS 



29 



which disappear towards the body whorl, was mentioned by 
Shepman for the much larger paratype. The prominent axial ribs 
on the subsutural slope are seen in Thiele's figure of the species. 
Judging from the published figures of B. trophonoidea, the 
species is rather variable, though it can be mentioned that the 
proportions of the shell (H/D and Hs/H) are rather constant in 
different specimens. Thiele's figure illustrates a somewhat more 
slender shell with better differentiated and more straight 
siphonal canal and much longer axial ribs, almost reaching the 
canal (they do not reach the whorl periphery in the holotype). 
Okutani published an illustration of a shell with a high and 
narrow spire. However, the variability of B. trophonoidea does 
not exceed that of the Atlantic representatives of the genus 
(Bouchet & Waren, 1980). 

Marshallena gracilispira Powell, 1969, described from Borneo 
and Philippines, 558-717 m, is probably a synonym of B. 
trophonoidea; from both original description and figure it is 
impossible to find any essential characters distinguishing the 
former species from the latter. However, this question cannot be 
resolved without an examination of type specimens and more 
material. 

Distribution. Gulf of Aden, Indonesia, and southern Japan, 
660-2000 m. The present record is the most western and most 
deep-sea locality. 

The following species were also mentioned from the John 
Murray Expedition bathyal samples by A. W.B.Powell (1964, 
1969) but not found in the material studied: 

Lucerapex denticulata (Thiele, 1925) - Powell, 1964, p. 286 (stn 
1 76, Gulf of Aden, 732 m; stn 1 84, Gulf of Aden, 1 270 m). 

Nihonia circumstricta (von Martens, 1901) - Powell, 1969, p. 334 
(stn 1 10, off Pemba Id., 333 m). 

LeucosyrinxjuliaThiele, 1925 -Powell, 1969, p. 338 (stn 34, Gulf 
of Aden, 1040 m). 

Typhlosyrinx vepallida (von Martens, 1902) - Powell, 1969, p. 
360 (stn 1 84, Gulf of Aden, 1 270 m). 

Marshallena philippinarum (Watson, 1882) - Powell, 1969, p. 
369-370 (three stations without numbers indicated, Gulf of 
Aden and off Pemba Id., 1061, 1022, and 802 m). 



DISCUSSION 

A total of 50 species of deep-sea conoidean gastropods were 
found in the JME collection. They belong to 3 families, 6 
subfamilies, 22 genera and 3 subgenera. Lower conoideans 
(families Drilliidae and Turridae) prevail in the material: 30 
species vs. 20 in Conidae. Among subfamilies, the most 
species-rich appeared to be Cochlespirinae (9 species) and 
Clathurellinae ( 1 1 species). 

The material studied seemingly does not represent well the 
fauna of the North- Western Indian Ocean; this is evidenced by 
low percentage of small species and juvenile individuals of larger 
species as compared, for example to East African bathyal fauna 
described by Thiele (1925). This is probably due to methods of 
collection. Additional evidence for this may be the very high 
share of lower conoideans, often respresented by large species, 
(60%) which far exceeds the respective values for East African 
fauna (about 41%, calculated from Thiele's (1925) list) and for 
other world-wide faunas (Sysoev, 1991). 

Nevertheless, some remarks on the JME collection can be 
made. The fauna has a typical bathyal appearance, with only few 



representatives of characteristic shallow-water (e.g. Drillia) and 
abyssal (e.g. Gymnobela (Theta) genera. 

The high percentage of new species in the material studied 
(about 1/3) indicates that the North- Western Indian Ocean is 
still insufficiently explored. This is additionally confirmed by the 
very low overlap with faunal lists obtained by other expeditions: 
for example, only 5 out of 59 bathyal species reported by Thiele 
(1925) for East Africa were found in the JME collection (plus 4 
species recorded by Powell - see above); the same is true for the 
bathyal fauna of Southern India collected by the 'Investigator' (7 
species found out of 37 reported by Winckworth, 1940). 

Most species were found at only one station, and thus the 
main areas covered by the JME investigations (i.e. Gulf of Aden, 
Zanzibar area, and Maldive Islands) have very few common 
species: 2 species (Comitas subsuturalis and Gemmula amabilis) 
were found in all three areas, 2 species (Gemmula bisinuata) and 
Borsonia symbiophora) - in the two first regions, and one species 
(Comitas erica) - in the two last. All species also found outside 
the region studied are apparently widely distributed in the 
Indo-Pacific. Of particular interest in this connection are the 
findings which greatly extend the geographic range of respective 
species: Horaiclavus splendidus previously known from Japan, 
Leucosyrinx claviforma from North-Western Australia, and 
Gymnobela daphnelloides from Hawaii. 



Acknowledgements. I am greatly indebted to Dr J.D.Taylor and Ms 
K. Way of The Natural History Museum, London for the loan of the 
material and providing relevant information. Dr J.D.Taylor and Dr 
PMordan of The Natural History Museum kindly edited the 
manuscript. Of great value were the taxonomic comments of an 
anonymous referee. I thank Dr Yu.I.Kantor of the A.N.Severtzov 
Institute of Problems of Evolution of Russian Academy of Sciences, 
Moscow, for his assistance. 



REFERENCES 



Abbott, R.T. & Dance, S.P. 1990. Compendium of seashells, 4th edition. 411 p. 

American Malacologists, Inc., Melbourne, Florida. 
Adams, A. 1867. Descriptions of new species of shells from Japan. Proceedings of 

the Zoological Society of London: 309-3 15. 
Alcock, A. & Anderson, A.R.S. 1898. Illustrations of the Zoology of the Royal 

Indian Marine Survey Ship Investigator, under the command of Commander 

T.H.Heming, R.N. Mollusca. Part 2. pi. 7-8. Calcutta. 
Alcock, A., Annandale N., & MacGilchrist, A.C. 1907. Illustrations of the Zoology 

of the Royal Indian Marine Survey Ship Investigator, under the command of 

Commander T.H.Heming, R.N. (Retired). Mollusca. Part 4. pi. 14-18. Calcutta. 
Alcock, A. & McArdle, A.F. 1 90 1 . Illustrations of the Zoology of the Royal Indian 

Marine Survey Ship Investigator, under the command of Commander 

T.H.Heming, R.N. Mollusca. Part 3. pi. 9-13. Calcutta. 
Annandale, N. & Stewart, F.N. 1910(1909]. Illustrations of the Zoology of the Royal 

Indian Marine Survey Ship Investigator, under the command of Captain 

W.G.Beauchamp. R. I. M. . Mollusca. Part 6. pi. 21-23. Calcutta. 
Bouchet, P. & Waren, A. 1980. Revision of the North-East Atlantic bathyal and 

abyssal Turridae (Mollusca, Gastropoda). Journal of Molluscan Studies 

supplement 8: 1-119. 
Cernohorsky, W.O. 1987. Taxonomic notes on some deep-water Turridae 

(Mollusca: Gastropoda) from the Malagasy Republic. Records of the Auckland 

Institute and Museum, 24: 123-1 34. 
Dall, W.H. 1895. Scientific results of explorations by the US Fish Commission 

streamer 'Albatross'. XXXIV. Report on Mollusca and Brachiopoda dredged in 

deep water, chiefly near the Hawaiian Islands, with illustrations of hitherto 

unfigured species from northwest America. Proceedings of the U.S. National 

Museum, 17(1032): 675-733. 
Gmelin, J.F. 1791. Caroli a Linne Systema naturae per regna tria Naturae. Editio 

decima tertia. 1(6), Vermes: 3021-3910. 
Gray, J.E. 1838. On some new species of quadrupeds and shells. Annals of natural 

History, 1(1): 27-30. 



30 



A.V. SYSOEV 



Habe, T. 1 970. Shells of the western Pacific in color. Vol. 2. Hoikusha, Osaka, 233 p. 
Kantor, Yu.I. & Sysoev, A.V. 1991 . Sexual dimorphism in the apertural notch of a 

new species of Gemmula (Gastropoda: Turridae). Journal of Molluscan Studies, 

57(2): 205-209. 
Kiener, L.C. 1839-40. Species general et iconographie des coquilles vivantes. 5, 

Genre Pleurotome. 84 p., 27 pis. J.B.Balliere et fils, Paris. 
Kilburn, R.N. 1971. Notes on some deep-water Volutidae, Turbinellidae and 

Turridae chiefly from off southern Mocambique and Natal with descriptions of 

two new species (Mollusca: Gastropoda). Annals of the Natal Museum, 21(1): 

123-133. 
1973. Notes on some benthic Mollusca from Natal and Mozambique with 

descriptions of new species and subspecies of Calliostoma, Solariella, Latiaxis, 

Babylonia, Fusinus, Bathytoma and Conus. Annals of the Natal Museum, 21(3): 

557-578. 

1983. Turridae (Mollusca: Gastropoda) of southern Africa and 

Mozambique. Part 1. Subfamily Turrinae. Annals of the Natal Museum, 25(2): 
549-585. 

1986. Turridae (Mollusca: Gastropoda) of southern Africa and Mozambique. 

Part 3. Subfamily Borsoniinae. Annals of the Natal Museum, 26: 417-470. 

1988. Turridae (Mollusca: Gastropoda) of southern Africa and 

Mozambique. Part 4. Subfamilies Drilliinae, Crassispirinae and Strictispirinae. 
Annals of the Natal Museum, 29(1): 167-320. 

1989. Notes on Plychobela and Brachytoma, with the description of a new 

species from Mozambique (Mollusca: Gastropoda: Turridae). Annals of the 
Natal Museum, 30: 185-196. 

Kosuge, S. 1988. Report on the family Turridae collected along the north western 

coast of Australia (Gastropoda) (3). Bulletin of the Institute of Malacology of 

Tokyo, 2(7): 118-123. 
1990. Report on the family Turridae collected along the north-western coast 

of Australia (Gastropoda) (4). Bulletin of the Institute of Malacology of Tokyo, 

2(9): 149-155. 
1992. Report on the family Turridae collected along the north-western coast 

of Australia (Gastropoda) (5). Bulletin of the Institute of Malacology of Tokyo, 

2(10): 162-174. 
Kuroda, T., Habe, T. & Oyama, K. 1971. The seashells of Sagami Bay. collected by 

His Majesty the Emperor of Japan. 714+429 p. Maruzen Co., Tokyo. 
Lamarck, J.B.P.A. de M. 1816. Tableau encyclopedique et methodique des trois 

regnes de la nature. Paris: Agasse. 1-16, pis 391-488. 
Martens, E. von 1901. Einige neue Meer-Conchylien von der deutsche 

Tiefsee-Expedition. Sitzungs-berichle der Gesellischaft naturforschender Freunde 

zu Berlin: 14-26. 
1902. Sitzungs-berichte der Gesellischaft naturforschender Freunde zu Berlin: 

19. 
1903 [1904]. Die beschalten Gastropoden der deutschen Tiefsee-Expedition 

1898-1899. A. Systematisch-geographischer Teil. Wissenschaftliche Ergebnisse 

der deutschen Tiefsee-Expedition aufdem Dampfer ' Valdivia' 1898-1899. Jena, G. 

Fischer, 7: 1-146. 
McLean, J.H. 1971. Family Turridae. pp. 686-766 In: Keen, A.M. Seashells of 

tropical West America; marine molluscs from Baja California to Peru. 2nd ed. 

Stanford, Stanford University Press. 
Melvill, J.C. 1910. Descriptions of twenty-nine species of marine Mollusca from 

the Persian Gulf, Gulf of Oman, and North Arabian Sea, mostly collected by 

Mr. FW.Townsend of the Indo-European Telegraph Service. Annals and 

Magazine of Natural History, ser. 8, 6: 1-17. 
Maxwell, P.A. 1988. Late Miocene deep-water Mollusca from the Stillwater 

Mudstone at Greymouth, Westland, New Zealand: paleoecology and 

systematica New Zealand Geological Survey Paleontological Bulletin, 55: 1-81 . 
Okutani, T. 1966. Archibenthal and abyssal Mollusca collected by the R./V 

Soyo-Maru from Japanese waters during 1964. Bulletin of the Tokai Regional 

Fisheries Research Laboratory, 46: 1-31 . 
Petuch, E. 1988. Neogene history of tropical American mollusks. 217 p. CERF, 

Charlottesville, Virginia. 
Powell, A.W.B. 1964. The family Turridae in the Indo-Pacific. Part 1. The 



subfamily Turrinae. Indo-Pacific Mollusca, 1(5): 227-346. 

1966. The molluscan families Speightiidae and Turridae. An evaluation the 

valid taxa, both Recent and fossil, with lists of characteristic species. Bulletin of 
the Auckland Institute and Museum, 5: 1 84 pp. 

1969. The family Turridae in the Indo-Pacific. Part 2. The subfamily 

Turriculinae. Indo-Pacific Mollusca. 2(10): 207^115. 

Schepman, M.M. 1913. Prosobranchia of the Siboga Expedition. Part 5. 

Toxoglossa. Siboga Expeditie Monograph, 49: 365-452. 
Shuto, T. 1965. Turrid gastropods from the Upper Pleistocene Moeshima Shell Bed 

(Molluscan palaeontology from Pleistocene formations in Kyushu -1). Memoirs 

of the Faculty of Science. Kyushu University, series D, Geology, 26(2): 143-207. 
1970. Taxonomical notes on the turrids of the Siboga-Collection originally 

described by M.M. Schepman, 1913 (Part I). Venus, 28(4): 161-178. 
1971. Taxonomical notes on the turrids of the Siboga-Collection originally 

described by M.M. Schepman. 1913 (Part III). Venus, 30(1): 5-22. 

1975. Notes on type species of some turrid genera based on the type 

specimens in the British Museum (N.H.). Venus, 33(4): 161-175. 

1983. New turrid taxa from the Australian waters. Memoirs of the Faculty of 

Science. Kyushu University, series D, Geology. 25( 1 ): 1-26. 
Smith, E.A. 1 894. Report upon some Mollusca dredged in the Bay of Bengal and 

the Arabian sea. Annals and Magazine of Natural History, ser. 6, 14: 1 57-174. 

1895. Report upon the Mollusca dredged in the Bay of Bengal and the 

Arabian sea during the season 1893-94. Annals and Magazine of Natural 
History, ser. 6, 16: 1-19. 

1896. Descriptions of new deep-sea Mollusca. Annals and Magazine of 

Natural History, ser. 6, 18: 367-375. 

1899. On Mollusca from the Bay of Bengal and the Arabian sea. Annals and 

Magazine of Natural History, ser. 7, 4: 237-251. 
1904. On Mollusca from the Bay of Bengal and the Arabian sea. Annals and 

Magazine of Natural History, ser. 7, 1 3: 453-473. 
1906. On Mollusca from the Bay of Bengal and the Arabian sea. Annals and 

Magazine of Natural History, ser. 1, 18: 157-185. 
Sturany, R. 1903. Gastropoden des Rothen Meeres. ExpeditionenS.M. Schijf'Pola' 

in das Rolhe Meer Nordliche und Sudliche Hafte 1895196-1897198. Zoologische 

ergebnisse. XXIII Berichte der Commission fur Oceanographische Forschungen. 

Wein. 75 pp, 7 pis. 
Sysoev, A.V. 1991. Preliminary analysis of the relationship between turrids 

(Gastropoda. Toxoglossa, Turridae) with different types of radular apparatus in 

various Recent and fossil faunas. Ruthenica. Russian Malacological Journal, 

1(1-2): 53-66. 
Sysoev, A.V. & Kantor, Yu.I. 1987. Three new species of the deep-sea molluscan 

genus Famelica (Gastropoda, Toxoglossa, Turridae). Zoologicheskij Zhurnal. 

56(8): 1255-1258 (In Russian). 
Taki, I. & Oyama, K. 1954. Matajiro Yokoyama's The Pliocene and later faunas 

from the Kwanto region in Japan. Paleontological Society of Japan Special 

Papers, 2: 1-68. 
Taylor, J.D., Kantor, Yu.I. & Sysoev, A.V. 1993. Foregut anatomy, feeding 

mechanisms, relationships and classification of the Conoidea (= Toxoglossa) 

(Gastropoda). Bulletin of the Natural History Museum, Zoology, 59(2): 125-170. 
Tesch, P. 1915. Jungtertiare und Quartare Mollusken von Timor. Teil I. 

Paldontologie von Timor, 5: 1-70. 
Thiele, J. 1925. Gastropoda der deutschen Tiefsee-Expedition. II Teil. 

Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition aufdem Dampfer 

■Valdivia 1898-1899. Jena, G.Fischer, 17(2): 1-348 (35-382) 
Watson, R.B. 1 88 1 . Mollusca of H . M .S. 'Challenger' expedition - part 9. Journal of 

the Linnean Society, 15: 413-455. 

1886. Report on the Scaphopoda and Gasteropoda collected by H.M.S. 

Challenger during the years 1 873-76. Reports of the Scientific Results of the 
voyage of H.M.S. Challenger, Zoology. 42: 1-756. 

Weinkauff, H.C. 1875. Die familie Pleurotomidae. Systematisches 
Conchylien-Cabinet , 4(3): 1-248, 43 pis. Nurenberg: Bauer & Raspe. 

Winckworth, R. 1940. A systematic list of the Investigator Mollusca. Proceedings 
of the Malacological Society of London, 24( 1 ): 1 9-29. 



Bull. nat. Hist. Mus. Lond. (Zool.) 62(1): 31-36 



Issued 27 June 1996 



Reassessment of 'Calcinus 9 astathes Stebbing, 

1924 

(Crustacea: Anomura: Paguridea: Diogenidae) 

patsy a. Mclaughlin 

Shannon Point Marine Center, 1900 Shannon Point Road, Anacortes, WA 98221-4042, U.S.A. 

CONTENTS 



Introduction 31 

Material and Methods 31 

Systematic Account 32 

Redescription 32 

Distribution 34 

Affinities 34 

Remarks 35 

Acknowledgements 35 

References 35 



Synopsis. A reexamination of the syntypes of 'Calcinus' astathes Stebbing, 1924 in the collection of The Natural History 
Museum, London. U.K., has shown that although subsequently assigned correctly to the genus Clibanarius, this taxon is 
not conspecific with C. virescens Krauss, 1 843, as was previously proposed at that time. Clibanarius astathes is redescribed 
and illustrated; a lectotype is designated. 



INTRODUCTION 



Stebbing (1924) described a new species of hermit crab from 
Delagoa Bay, South Africa, which he mistakenly assigned to 
Calcinus Dana, 1851. The fact that this species clearly should 
have been placed in Clibanarius Dana, 1852 was recognized by 
Barnard (1947); however, at that time he made only the notation 
that four specimens returned to the South African Museum by 
Stebbing were indistinguishable from Clibanarius virescens 
(Krauss, 1843). Barnard (1950) went on to define those four 
specimens further by commenting, 'I would not have ventured to 
dispute the identity of Stebbing's Calcinus astathes if there had 
not been four specimens returned to the Museum bearing 
Stebbing's autographic label (the largest and type specimen 
probably retained by him, or perhaps now transferred to the 
British Museum). These four specimens are obviously C. 
virescens. The dactyl of the 3rd leg is not longer than 6th joint, 
and has the characteristic shape'. 

Stebbing's (1924) original description of Calcinus astathes 
specified only a 'group' of specimens collected from Delagoa 
Bay by K.H. Barnard on October 12, 1912. Five specimens, 
listed as syntypes of this taxon are part of the Stebbing 
Collection donated to The Natural History Museum (NHM) 
[formerly British Museum (Natural History)] by Barnard, 
although only three initially were listed in the Museum registry 
(NHM 1928.12.1.264-266). Four specimens, dry, and in poor 
condition, remain in the collection of the South African 
Museum (SAM) catalogued under the original number (A2121) 



listed by Stebbing (1924). Barnard's hand written label 
accompanying the dry specimens reads 'A2121 . Delagoa Bay 4 
spec, returned by Stebbing labelled (sic) as 'Calcinus astathes' 
also 1 with parasites see Stebbing. 1920'. Barnard's label 
reference to the parasitized specimen must refer to a specimen 
without chelipeds included by Stebbing (1920), with others 
(NHM 1928.12.1.267-268, SAM A3270 A2120), identified as 
Calcinus laevimanus (Randall, 1840), the former specimen 
similarly placed in synonymy with C. virescens by Barnard 
(1950: 435). No specimens of Calcinus or Clibanarius astathes 
were listed among the type collection in the South African 
museum by Kensley (1974), and none have been found on a 
recent search (Ms. L. Hoenson, pers. comm.). 

Despite some confusion in the initial NHM registry entry for 
' Calcinus' astathes, at least two points are clear. These five 
specimens labeled as syntypes of Stebbing's (1924) species all 
belong to the same taxon; none are conspecific with Clibanarius 
virescens. The four specimens remaining in the SAM collection 
truly do represent C virescens. The largest of the NHM 
specimens [and the one presumably unaccounted for by Barnard 
(1950)] is herein designated as the lectotype of 'Calcinus' 
astathes. 



MATERIAL AND METHODS 



In addition to the syntypes, and Barnard's four specimens of 
Clibanarius virescens, comparative material of a 



©The Natural History Museum, 1996 



32 



pa. Mclaughlin 



morphologically similar Clibanarius species, C. longitarsus (De 
Haan, 1849) from Durban Bay, S.A., has been examined. One 
measurement, shield length (SL), measured from the tip of the 
rostrum to the midpoint of the cervical groove provides an 
indication of specimen size. The type material of C. astathes has 
been returned to the Natural History Museum, and Barnard's C. 
virescens to the South African Museum. Two specimens of C. 
longitarsus used in the comparison also have been deposited in 
The Natural History Museum (NHM 1995.163-164), the 
remaining have been retained in the author's personal collection. 



SYSTEMATIC ACCOUNT 



Clibanarius astathes (Stebbing, 1924) 

Figs 1 , 2 

Calcinus astathes Stebbing, 1924: 239, pi. 2 (CXVII of 
continuing series). Clibanarius virescens: Barnard, 1947: 376 (in 
part); 1950: 435 (in part). Lectotype (herein selected). Female 
(SL - 7.3 mm), NHM reg. 1928.12.1.264 Paralectotypes. Two 
females (SL = 6.3, 6.3 mm), 2 males (SL = 5.8, 6.9), NHM reg. 
1928.12.1.265-266. 

Type locality. Delagoa Bay (25°50'S, 32°50'E), 
'Mosambique', Ethiopian region, Indian Ocean, 1912. 

Diagnosis. Rostrum with simple or bifid termination. Ocular 
peduncles 0.66-0.75 length of shield; little if at all overreached 
by antennular peduncles; antennal peduncles not reaching to 
bases of corneae. Ocular acicles with acute, simple or bifid 
termination. Basal segment of antennular peduncle with 1 or 2 
small spines. 

Chelipeds subequal, right somewhat larger. Right cheliped 
with dactyl equal to or slightly longer than palm; dorsomesial 
margin and dorsal surface with rows of corneous-tipped spines. 
Dorsomesial margin of palm with row of low spines and tufts of 
long setae, second adjacent row of spines and one large tubercle 
at proximal margin, dorsal surface sloping, with few spines 
distally and on proximal portion of fixed finger. Dorsomesial 
distal angle of carpus with acute spine, low protuberances and 
tufts of setae on dorsomesial margin. Merus with few low, 
somewhat spinulose protuberances on ventromesial margin; 
ventrolateral distal angle with three acute spines. Left cheliped 
with few more spinules on dorsal surface of palm. Carpus with 
two strong corneous-tipped spines on dorsomesial margin and 
blunt protuberance in line with tubercle on proximal margin of 
palm. Ventromesial margin of merus with row of few small 
spines or two small spines distally; lateral face with two spines at 
ventrolateral distal angle. 

Ambulatory legs similar from left to right. Dactyls 
approximately 1.5 times longer than propodi; dorsal surfaces 
flattened; ventral margins each with row of minute corneous 
spinules ( 1 0- 1 3 in distal half and one or two proximally). Lateral 
faces of propodi each with very strongly developed dorsolateral 
margin; dorsal surfaces somewhat flattened; ventrolateral distal 
angles each with one or two spines. Carpi each with spine at 
dorsodistal angle. Meri each with acute spine at ventrolateral 
distal angle. Sternite of third pereopods with subrectangular 
anterior lobe, anterior margin slightly rounded. Fourth 
pereopods each with acute spine at dorsodistal margin of 
carpus. 

Telson with slightly asymmetrical posterior lobes, separated 



by small median cleft; terminal margins each with three to five 
small spines, larger on left. 



REDESCRIPTION 

Shield longer than broad; anterior margin between rostrum and 
lateral projections straight or very faintly concave; anterolateral 
margins sloping; posterior margin roundly truncate. Dorsal 
surface of shield with scattered setae and distinct 'Y'-shaped 
suture medianly in posterior portion. Rostrum triangular, 
terminating acutely or minutely bifid, overreaching lateral 
projections. Lateral projections broadly rounded or obtusely 
triangular, with terminal spinule or blunt small projection. 

Ocular peduncles slender, 0.66-0.75 length of shield, with 
scattered setae dorsally and mesially; corneae not dilated. 
Ocular acicles narrowly triangular, dorsally rounded (convex), 
with simple or bifid acute termination; separated by less than 
basal width of one acicle and tending to become approximate 
distally. 

Antennular peduncles overreaching distal margin of corneae 
little if at all. Ultimate and penultimate segments with scattered 
setae. Basal segment with one or two very small spinules on 
ventrolateral distal margin. 

Antennal peduncles reaching approximately to bases of 
corneae; with supernumerary segmentation. Basal segment 
without spine on laterodistal margin, but with acute spine on 
ventrodistal margin laterally. Second segment with dorsolateral 
distal angle produced, with small terminal spine, dorsomesial 
distal angle rounded. Third segment with small spine on 
ventrodistal margin. Fourth and fifth segments with scattered 
setae. Antennal acicle reaching almost to distal margin of fourth 
peduncular segment, triangular, armed on mesial margin with 
one to three spines and tufts of setae, terminating in acute spine. 
Antennal flagellum overreaching chelipeds, and approximately 
as long as ambulatory legs; each article with two or three minute 
bristles. 

Chelipeds subequal, right somewhat larger. Right cheliped 
with dactyl equal to or slightly longer than palm; dorsomesial 
margin with row of corneous-tipped spines, dorsal surface with 
two rows of appreciably stronger, corneous-tipped spines; 
surfaces all with numerous tufts of moderately long setae; 
cutting edge with two prominent calcareous teeth in proximal 
half and broad terminal corneous hoof-shaped claw. Palm 
slightly longer than carpus; dorsomesial margin with row of low 
spines and tufts of long setae, second adjacent row of spines and 
one large tubercle at proximal margin, dorsal surface sloping, 
with no delimitation of dorsolateral margin, surface with few 
spines distally and on proximal portion of fixed finger, also with 
tufts of long setae; fixed finger with two rows of small spines on 
dorsal surface, all surfaces with tufts of long setae; cutting edge 
with three calcareous teeth in proximal half, distal-most 
strongest; terminating in corneous hoof-shaped claw. Carpus 
slightly more than half length of merus; dorsomesial distal angle 
with acute spine, low protuberances and tufts of setae on 
dorsomesial margin; dorsal surface with indications of points of 
original tufts of setae (no longer present), dorsolateral margin 
not delimited; mesial face with few scattered tufts of setae. 
Merus subtriangular; dorsal margin with tufts of setae; 
ventromesial margin with few low, somewhat spinulose 
protuberances and tufts of setae; ventrolateral distal angle with 
three acute spines, ventrolateral margin proximally and lateral 






REASSESSMENT OF CALCINUS' ASTATHES 



33 




Fig. 1 Clibanarius astathes (Stebbing, 1924), A-E, female lectotype (7.3 mm); F, male paralectotype (6.9 mm). A, shield and cephalic appendages; B, 
right second antennal peduncular segment enlarged; C, carpus and chela of left cheliped (lateral view); D, third left pereopod (lateral view); E, dac- 
tyl of third left pereopod (mesial view); F, telson. Scales equal 5.0 mm (A, C-E) and 3.0 mm (B, F). 



34 



p.a. Mclaughlin 



face ventrally with low protuberances and tufts of setae. Ischium 
with few low spinules on ventromesial margin. Left cheliped 
with few more spinules on dorsal surface of palm; carpus with 
two strong corneous-tipped spines on dorsomesial margin and 
blunt protuberance in line with tubercle on proximal margin of 
palm; ventromesial margin of merus with row of few small 
spines or two small spines distally and two, plus low 
protuberance, medially, with additional protuberance 
proximally; lateral face with two spines at ventrolateral distal 
angle and low protuberances on ventrolateral margin. 

Ambulatory legs similar from left to right. Dactyls 
approximately 1.5 times longer than propodi; in dorsal view 
straight, in lateral view slightly curved; dorsal surfaces flattened, 
and with rows of tufts of stiff setae, lateral faces each with 
longitudinal row of tufts of stiff setae; ventral margins with row 
of tufts of stiff setae and row of minute corneous spinules, 14 
(spaced distally to proximally 8,5,1) on left third (lectotype), 
right third with 10, left second with 13, all in distal half; 
paralectotypes usually with 10-13 spinules in distal half and one 
or two proximally. Propodi slightly less than twice length of 
carpi; lateral faces each with distinctly developed dorsolateral 
margin and tufts of setae; dorsal surfaces somewhat flattened 
and with numerous tufts of setae; ventrolateral distal angles 
each with one or two spines, ventral margins with tufts of setae. 
Carpi 0.66 to 0.90 length of meri; each with spine at dorsodistal 
margin and tufts of setae on dorsal and lateral faces. Meri with 
tufts of setae dorsally and ventrally; ventrolateral distal angles 
each with one acute spine and ventral margins of second also 
with low protuberances. Ischia with tufts of setae on ventral 
margins. Fourth pereopods each with acute spine at dorsodistal 
margin of carpi. Sternite of third pereopods with 
subrectangular anterior lobe, anterior margin slightly rounded. 
Uropods asymmetrical. Telson with transverse suture; 
posterior lobes slightly asymmetrical, separated by small 
median cleft; terminal margins each with three to five small 
spines, larger on left. 

Colour. Unknown. 

Distribution. At present recognized only from the type 
locality, Delagoa Bay, South Africa. 



Affinities. Clibanarius astathes shares with C. padavensis De 
Man, 1888, dactyls of the ambulatory legs that are longer than 
the propodi. Barnard (1926) listed the latter species from 
Delagoa Bay, and subsequently (Barnard, 1950) gave a brief 
diagnosis of that species. The ratio of cornea diameter to ocular 
peduncle length cited by Barnard is less than that given by De 
Man ( 1 888) in his original description, or later by Alcock (1905), 
and the illustrated propodus of the left third pereopod (Barnard, 
1950: fig. 80d) is appreciably shorter and stouter than that 
described for C. padavensis. However, if the colour patterns 
described by Barnard actually were taken from his specimens, 
one must assume that his identification was correct, and that C. 
astathes and C. padavensis exist sympatrically in the Delagoa 
area. 

I have not had the opportunity to examine Barnard's (1950) 
specimen(s) of C. padavensis, nor other specimens of this 
species; however, De Man's (1888) very detailed description 
points to several characters that would distinguish C. padavensis 
from C. astathes in the absence of colour. These include longer 
and more slender ocular peduncles; multispinose ocular acicles; 
longer antennal acicles, which reach beyond the proximal 
margins of the penultimate peduncular segments; longer and 
more slender chelae; and subcylindrical propodi of the 
ambulatory legs. 

Clibanarius astathes also bears a very strong resemblance to 
C. longitarsus. Clibanarius astathes differs from the Durban Bay 
population of C. longitarsus (12 males, 3 females, SL = 1.9-10.4 
mm) that I have examined, in having: 1) fewer spines on the 
ventrolateral distal margin of the antennular peduncle (one or 
two, as opposed to three to six in C. longitarsus); 2) two strong 
corneous-tipped spines on the dorsomesial margin of the carpus 
of the left cheliped (only one was observed in numerous 
specimens of C. longitarsus of varying sizes); 3) strongly 
delineated dorsomesial propodal margin on the third pereopod 
(rounded or very faintly ridged in C. longitarsus); 4) roundly 
rectangular anterior lobe on the sternite of the third pereopods 
(this lobe is subquadrate, and often with a central blister-like 
protuberance in C. longitarsus). Other characters, such as the 
rows of spines on the fixed finger of the left cheliped, the 
armature of the ventral margins of the meri of the chelipeds, the 







Fig. 2 Clibanarius astathes (Stebbing, 1924), male paralectotype (6.9 mm). A, ieft cheliped; B, right cheliped. Scale equals 12 mm. 



REASSESSMENT OF 'CALCINUS' ASTATHES 



35 



presence of a spine at the ventrolateral distal angle of the merus 
of the third right and left pereopods, the number and spacing of 
the corneous spinules on the ventral margins of the ambulatory 
dactyls, and the spination of the terminal margins of the telson 
appear quite variable in C longitarsus. Too few specimens of C. 
astathes are known for any evaluation of morphological 
variation. 

Remarks. Stebbing's (1924) comments on the genus Calcinus 
emphasized as did his earlier remarks (Stebbing, 1914), the 
development of the maxillipeds, which seems to suggest that he 
was not really familiar with the overall morphology of Calcinus 
species. Finding similar maxillipedal development in his new 
taxon, Stebbing (1924) assigned astathes to this genus, with 
apparent disregard for the numerous characters which set 
Clibanarius apart from Calcinus. 

It also would appear that the telson of the specimen he 
described and illustrated (Stebbing, 1924: 240, pi. 2T) was not 
closely examined, as it was characterized and depicted as being a 
simple plate with a smoothly rounded terminal margin. In 
actuality, the telson has a slight transverse suture dividing it into 
anterior and posterior lobes; the posterior lobes are separated by 
a small median cleft, and the terminal margins each have a few 
distinct spines. 

Barnard (1950) was correct to transfer Stebbing's ( 1 924) taxon 
to Clibanarius, although this transfer was obscured by his 
placement of the species in synonymy with C. virescens. Calcinus 
astathes Stebbing, 1924 was still listed by Gordan (1956) in her 
comprehensive tabulation of hermit crab species, and more 
recently was included by Morgan ( 1991 ) in his world-wide listing 
of known Calcinus species. Barnard's (1950) decision regarding 
the conspecificity of Clibanarius astathes and C. virescens 
appears to have been based on specimens incorrectly labeled by 
Stebbing, and not on the actual type material of C astathes. Not 
only the fact that the Natal-Mozambique areas of South Africa 
are type localities of both species, but also the inadequacy of 
Stebbing's (1924) original description and illustrations, 
undoubtedly account for Barnard's (1950) synonymy having 
been accepted, without question, by subsequent carcinologists. 

One of the principal characters upon which Barnard (1950) 
based his identification of C. virescens was the shortness of the 
dactyls of the third pereopods in relation to the propodi, 
although, as pointed out by Lewinsohn (1982), this character 
was not mentioned in Krauss' (1843) original description of the 
species. Not only Barnard (1950), but Fize and Serene (1955), 
and Gherardi and McLaughlin (1994) reported that the dactyls 
of their specimens were shorter than the propodi; however, 
Buitendijk (1937), Miyake (1978), and Lewinsohn (1982) 
described the dactyls and propodi as being equal in length. 
Rahayu and Forest (1993) found the dactyls longer than the 
propodi in small specimens, but shorter in large specimens. 
Stebbing (1924), like Krauss (1843), made no mention of the 
length ratios of the dactyls and propodi of the ambulatory legs 
of C. astathes. 

Despite the variations in this major diagnostic character 
observed in C. virescens, C. astathes sensu stricto differs 
markedly from Krauss' (1843) taxon. The dactyls of the 
ambulatory legs of C. astathes are approximately half again the 
length of the propodi. Additionally, the ventral margins of the 
ambulatory dactyls are armed with 10 to 14 tiny corneous 
spinules in C. astathes, in contrast to the five to eight strong 
corneous spines seen in C virescens. As previously indicated, C. 
astathes bears a far greater similarity to that group of 



Clibanarius species characterized by long pereopodal dactyls 
and very short antennal acicles. 



Acknowledgements. I am indebted to Paul Clark, The Natural 
History Museum, London, for the loan of Stebbing's syntypic material, 
and to Barbara Cook and Liz Hoenson, South African Museum, Cape 
Town, for providing the specimens upon which Barnard based his 
synonymy. The photographs are to work of E.J. McGeorge. This a 
scientific contribution from the Shannon Point Marine Center, Western 
Washington University. 



REFERENCES 



Alcock, A. 1905. Anomura. Fast: I. Pagurides. - Catalogue of the Indian decapod 

Crustacea in the collections of the Indian Museum 2: 1-197. Calcutta. Indian 

Museum. 
Barnard, K. H. 1926. Report on a collection of Crustacea from Portuguese East 

Africa. Transactions of the Royal Society of South Africa 13(2): 1 19-129. 
1947. Descriptions of new species of South African decapod Crustacea, with 

notes on synonymy and new records. The Annals and Magazine of Natural 

History (II) 13:361-392. 
1950. Descriptive catalogue of South African decapod Crustacea (crabs and 

shrimps). Annals of the South African Museum 38: 1 -837. 
Buitendijk, A. M. 1937. Biological results of the Snellius expedition. IV. The 

Paguridea of the Snellius Expedition. Temminckia 2: 251-280. 
Dana, J.D. 1851. Conspectus crustaceorum quae in orbis terrarum 

circumnavigatione, Carolo Wilkes e classe reipublicae foederatae duce, lexit et 

descripsit. (Preprint (rom)Proceedings of the Academy of Natural Sciences. 

Philadelphia 5: 267-272. 
1852. Conspectus crustaceorum, etc.. Conspectus of the Crustacea of the 

Exploring Expedition under Capt. Wilkes, U.S.N., including the Paguridea, 

continued, the Megalopidea, and the Macroura. Paguridea, continued, and 

subtribe Megalopidea. (Preprint from) Proceedings of the Academy of Natural 

Sciences. Philadelphia 6: 6-28 ( 1 854). 
Fize, A. & Serene, R. 1955. Les Pagures du Vietnam. Institut Oceanographiquc 

Nhatrang Note 45: ix, 1-228. 
Gherardi, F. & McLaughlin, P. A. 1994. Shallow-water hermit crabs (Crustacea: 

Decapoda: Anomura: Paguridea) from Mauritius and Rodrigues Islands, with 

the description of a new species of Calcinus. Raffles Bulletin of Zoology 42(3): 

613-656. 
Gordan, J. 1956. A bibliography of pagurid crabs, exclusive of Alcock, 1905. 

Bulletin of the American Museum of Natural History 108: 253-352. 
Haan, W., De. 1833-1850. Crustacea. In: P.F. von Siebold, Fauna Japonica wive 

Descriptio animalium. quae in ilinere per Japoniam.jussu et auspicilis super iorum, 

qui summum in India Batava Imperium lenent. suscepto. annis 1823-1830 collegit, 

notis. ohservationibus et adumhrationibus illustravit: 4: ix-xvi, vii-xvii, i-xxxi, 

1-244, pis. 1-55. Lugdunum Batavorum. 
Krauss, F. 1843. Die Sudafrikanischen Crustaceen. Eine Zusammenstellung alter 

bekannten Malacostraca. Bemerkungen uber deren Lebensweise und 

geographische Verbreitung. nebst Beschreibung und Abbildung mehrerer neuen 

Arlen. 68 pp. E. Schweizerbart'sche Verlagsvuchhandlung, Stuttgart. 
Kensley, B. 1974. Type specimens of Decapoda (Crustacea) in the collections of the 

South African Museum. Annals of the South African Museum 66(4): 55-77. 
Lewinsohn, Ch. 1982. Researches on the coast of Somalia. The shore and the dune 

of Sar Uanle. 33. Diogenidae, Paguridae and Coenobitidae (Crustacea 

Decapoda Paguridea). Monitore Zoologico Italiano. n.s. supplement 16: 33-68. 
Man, J. G., De. 1888. Report on the Podophthalmous Crustacea of the Mergui 

Archipelago, collected for the Trustees of the Indian Museum, Calcutta, by Dr. 

John Anderson, F.R.S., Superintendent of the Museum, parts IV and V. Journal 

of the Linnean Society. London 22:1 77-246. 
Miyake, S. 1978. The crustacean Anomura of Sagami Bay: 1-200 (English), 1-161 

(Japanese). Hoikusha Publishing Co., Tokyo. 
Morgan, G.J. 1991. A review of the hermit crab genus Calcinus Dana (Crustacea: 

Decapoda: Diogenidae) from Australia, with descriptions of two new species. 

Invertebrate Taxonomy 5: 869-913. 
Rahayu, D.L. & Forest, J. 1993. Le genre Clibanarius (Crustacea, Decapoda, 

Diogenidae) en Indonesie, avec la description de six especes nouvelles. Bulletin du 

Museum National d'Histoire Naturelle. Par is [\992] (4) 14(A)(2): 745-779. 
Randall, J.W. 1840. Catalogue of the Crustacea brought by Thomas Nuttall and 

J.K. Townsend, from the west coast of North America and the Sandwich Islands, 

with descriptions of such species as are apparently new . . . Journal of the 

Academy of Natural Sciences of Philadelphia 8: 1 06- 1 47. 



36 

Stebbing, T.R.R. 1914. Stalk-eyed Crustacea Malacostraca of the Scottish 

National Antarctic Expedition. Transaction of the Royal Society of Edinburgh, 

50(2): 253-307 (issued separately June 4, 1 9 14). 
1920. South African Crustacea (Part X of S. A. Crustacea, for the Marine 

Investigations in South Africa). Annals of the South African Museum. 17(4): 

231-272. 



pa. Mclaughlin 

1924. South African Crustacea (Part XII of S. A. Crustacea, for the Marine 

Investigations in South Africa). Annals of the South African Museum 19: 
235-250. 



Bull. nat. Hist. Mus. Lond. (Zool.)62(l): 37-39 



Issued 27 June 1996 



On a new species of Ophidiaster 
(Echinodermata: Asteroidea) from southern 
China 

YULIN LIAO 

Institute of Oceanology, Academia Sinica, 7 Nanhai Road, Qingdao, RR. China 

AILSA M.CLARK 

Formerly of The Natural History Museum, Cromwell Road, London SW7 5BD, UK 

Synopsis. A new species, Ophidiaster multispinus, from southern Chinese waters, is described. This was previously 
recorded by A.M . Clark ( 1 982) and by Liao & Clark (in press) as Ophidiaster armatus Koehler, 1910 but both authors now 
believe that Chinese specimens are specifically distinct. 



SYSTEMATIC DESCRIPTION 



Family OPHIDIASTERIDAE 
Genus Ophidiaster L. Agassiz, 1835 

Ophidiaster multispinus sp. nov. 

Fig. 1, pi. 1 

Ophidiaster armatus: A.M. Clark, 1982:487, 490; Liao & Clark 
(in press): .[Non Ophidiaster armatus Koehler, 1910]. 

HOLOTYPE. IOAS:E1070, Hainan Strait, southern China 
(20°15'N, 110°15'E), 55 m, rocky, collected July 10, 1960; 
paratypes IOAS-E-1071, 4 specimens from Xiamen (Amoy), 
Fujian Province, 1975. 

Description. R (major radius) 55-70 mm; r (minor radius) 10 
mm, br (arm breadth basally) 1 1 mm. Disc small, arms five, 
unequal, cylindrical, only tapering slightly in the distal third to 
rounded tips. Abactinal plates large, more or less triangular in 



shape, slightly convex, covered with numerous coarse granules, 
density in the central part of the plates 12-18/mm 2 . Abactinal 
plates on dorsal surface of arms arranged in three regular 
longitudinal series, together with the two marginal series each 
side forming seven regular longitudinal series as well as 
transverse ones. Papular areas distinctly arranged in eight 
longitudinal series but the lowest on each side with only a few 
pores in each area. No pedicellariae detected. All the marginals, 
except the basal three to five, armed with a fairly conspicuous 
short blunt spine, these forming a longitudinal series but missing 
on occasional plates, leaving small gaps in the sequence. Some 
abactinal plates near the arm tips also bearing one or two small 
spines but these are not at all conspicuous. 

Adambulacral plates with two furrow spines, those of 
consecutive plates not separated by granules on the vertical faces 
of the furrows. A large cone-like subambulacral spine on each 
plate, set back from the furrow within the general granulation. 
The most proximal six to eleven adambulacral plates with a 
smaller supplemental series of subambulacral spines interposed 
between the furrow spines and the main subambulacral ones. 




Fig. 1 Ophidiaster multispinus sp. nov., Holotype. Proximal portion of actinal surface showing: (a) furrow spines; (b) supplemental subambulacral 
spines; (c) subambulacral spines; (d) actinal spines. The mouth is towards the right. The scale bar = 2 mm. 



l©The Natural History Museum, 1996 



38 



Y. LIAO AND A.M. CLARK 





Plate 1 Ophidiaster multispinus sp. nov. "Holotype: (a) dorsal view; (b) ventral view. 



NEW SPECIES OF OPHIDIASTER 



39 



Four or five actinal plates in each intermediate area, armed with 
single spines corresponding to the subambulacral spines (Fig. 1 ). 
Colour (in the dried state) yellowish with faint darker bands. 

In addition to the holotype, there are four paratypes from 
Amoy, Fujian Province, far to the east of Hainan Strait. These 
are much larger than the holotype (which was selected for better 
comparison with the type material of Ophidiaster armatus) and 
have R 95, 105, 108 and 115 mm. Three of them have 
pedicellariae on some abactinal plates. 

Four other large specimens (R c. 1 30 mm), from the east side 
of Hong Kong Island in c. 15 m, studied by A.M.C., show some 
variation in the extent of the marginal spines, which are most 
numerous on the inferomarginals and may extend almost to the 
interradius. 

Remarks. This new species was previously referred to 
Ophidiaster armatus Koehler, 1910 by A.M. Clark (1982) and by 
Liao & Clark in 'The echinoderms of southern China' (in press), 
both references commenting on the unusually large size. During 
a visit to the Senckenberg Museum by the senior author in 1993, 
a direct comparison was made between the small Chinese 
specimen and the three syntypes of O. armatus from the Aru 
Islands, Indonesia, of similar size. As a result of this 
comparison, we conclude that the Chinese specimens are 
specifically distinct, the differences being evident as shown in 
Table 1. 

The maximum size for Ophidiaster armatus, R 66 mm, was 
observed by H.L. Clark (1938) in material from Queensland, 
Australia. 

It is noteworthy that all three syntypes of O. armatus do have 
some abactinal pedicellariae which Koehler seems to have 
overlooked, however, the occurrence of pedicellariae is rarely 
regarded as a character of specific weight in this family. 

The presence of spines on plates other than the 
adambulacrals, separates Ophidiaster armatus and O. 
multispinus from the remaining species of the genus Ophidiaster. 



Table 1 Comparison between the two species mentioned. 

Ophidiaster multispinus sp. nov. O. armatus Koehler 

Maximum R 130 mm. Maximum R 66 mm. 

Arms stout, br 1 1 mm at R 60 mm. Arms slender, br 9 mm at R 60 mm. 

Marginal spines fairly conspicuous Marginal spines not at all 
and more extensive, absent only conspicuous, only present 
from the first 3-5 plates towards the arm tips 

Proximal adambulacrals with two Subambulacral spine series 
series of subambulacral spines single throughout 



4 or 5 spinose actinal plates in 
intermediate area 



No spinose actinal plates 



However, in our opinion, this character is not of sufficient 
importance to justify a generic distinction, especially as the 
lesser development in O. armatus provides an intermediate 
condition. 



REFERENCES 



Clark, A.M. 1982. Echinoderms of Hong Kong, pp. 485-500. In: B.S. Morton & 

C.K. Tseng (Eds) The marine flora and fauna of Hong Kong and southern China. 

(Hong Kong, Hong Kong University Press). 
Clark, A.M. & Rowe, F.W.E. 1971. Monograph of shallow-water Indo-West Pacific 

echinoderms. London: British Museum (Natural History), ix + 238 pp. 
Clark, H.L. 1938. Echinoderms from Australia. Memoirs of the Museum of 

Comparative Zoology. Harvard 55: viii + 596 pp. 
Koehler, R. 1910. Asteries et Ophiures des iles Aru et Kei. Abhandlungen 

Senckenbergischen Naturforschen dem Gesellschaft, 33(3): 265-295. 
Liao, Y. & Clark, A.M. (in press). The echinoderms of southern China. 



Bull. nat. Hist. Mus. Lond. (Zool.) 62(1): 41-70 



Issued 27 June 1996 



The life cycle of Paracyclops fimbriatus (Fischer, 
1853) (Copepoda, Cyclopoida) 

S. KARAYTUG 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5 BD, UK, and School 
of Biological Sciences, Queen Mary and Westfield College, Mile End Road, London El 4NS, UK 

G. A. BOXSHALL 

Department of Zoology, The Natural History Museum, Cromwell Road, London SW7 5 BD, UK 

CONTENTS 



Introduction 41 

Methods 41 

Description of Naupliar Stages 42 

Description of Copepodid Stages 48 

Discussion 67 

Acknowledgements 69 

References 70 



Synopsis. The complete life cycle of Paracyclops fimbriatus (Fischer, 1 853) (Copepoda, Cyclopoida) is redescribed based 
on cultured material. All 6 nauplius and 5 copepodid stages plus the adults are described in detail. Scanning electron 
microscopy is used to elucidate the fine structure of some appendages, especially the male antennule, and to study body 
ornamentation patterns. Sexual dimorphism is first apparent at the copepodid IV stage which has a more robust antennule 
in the male than in the female. 



INTRODUCTION 



Paracyclops fimbriatus has been reported from a wide range of 
freshwater habitats and the geographical records summarised by 
Dussart & Defaye (1985) indicate a cosmopolitan distribution. 
It has also been regularly recorded from subterranean waters 
(ltd, 1954;Pesce&Galassi, 1987) and from waters with a salinity 
of 31%o(L6ffler, 1961). Early reports of this species at great 
depths in Lake Baikal (Mazepova, 1962) refer to the newly 
recognized R baikalensis Mazepova, 1978 (Mazepova, 1978). 

The genus Paracyclops comprises 24 species and subspecies 
(Dussart & Defaye, 1985; Reid, 1987a, 1987b; Strayer, 1988). 
The type species, P. fimbriatus, was originally described by 
Fischer (1853), using material from St-Petersburg in Russia. 
This taxon has been partially redescribed on numerous 
occasions (for example Kiefer, 1929; Gurney, 1933; Lindberg, 
1941; Dussart, 1969; Einsle, 1971). Some descriptions of 
developmental stages have also been published; one of the most 
accurate being that of Gurney (1933) but even that gives 
insufficient data to characterise the copepodid stages. Other 
descriptions have concentrated primarily on the naupliar instars 
(Gurney, 1933; Dukina, 1956; Ewers, 1930), but these all lack 
detail by modern standards. Although the general morphology 
of cyclopoid naupliar stages has been known for a long time it is 
only recently that detailed naupliar descriptions of freshwater 
cyclopoid copepods have been published (Dahms & Fernando 
1992, 1993, 1994). Systematic studies on naupliar stages can 



provide important information on the phylogenetic 
relationships among copepods (Dahms, 1991a, b). 

In the present paper P. fimbriatus has been redescribed in 
detail, including all its developmental stages, in order to provide 
a base for comparison with other Paracyclops species. 



METHODS 

Adult females of P. fimbriatus were collected by plankton net 
from Regent's Park pond (London) on 29.5.1994. Ovigerous 
females were kept in plastic dishes filled with fresh water filtered 
through a 35 um-mesh net. Water was changed at 2 day intervals 
until the eggs hatched and the nauplius phase was completed. 
The adult females were removed after hatching of their egg sacs. 
Copepodid stages were placed in an aquarium tank in a constant 
temperature room. 

The nauplii were examined as whole mounts. Broken 
glass-fibres were used to prevent the nauplii from being 
compressed by the coverslip and to facilitate rotation which 
allowed viewing from all sides. 

Copepodids were dissected in lactophenol. All drawings were 
made with the aid of a camera lucida using Nomarski 
differential interference contrast. All measurements were made 
with an ocular micrometer. Body lengths were measured from 
the anterior to the posterior end of the body in nauplii, and from 
the base of the rostrum to the posterior edge of the caudal rami 



©The Natural History Museum, 1996 



42 



S. KARAYTUG AND G.A. BOXSHALL 



in copepodids. Body width is given as the widest part of the 
nauplius body or the copepodid cephalothorax. In the spine and 
seta formula of the swimming legs Roman numerals and Arabic 
numerals are used for spines and setae respectively. 

When necessary scanning electron microscopy (SEM) was 
used to study fine details. Material for SEM was prepared by 
cleaning specimens in an aqueous solution (one drop in 100 ml 
of distilled water) of the surfactant detergent RBS pF for 30 min 
and subsequently sonicating in an ultrasonic cell disrupter for 7 
s. Cleaned specimens were washed in 3 changes of distilled water 
for 5 minutes, dehydrated through a graded acetone series, 
critical point dried, mounted on aluminium stubs, sputter 
coated with gold and viewed under an Hitachi S-800 scanning 
elctron microscope. 



RESULTS 



Description of Naupliar Stages 

Nauplius I 

Body length : mean ± standard deviation = 1 1 8 ± 2.68 urn (range 
1 1 3 to 1 22 um, n=20), mean body width 75 ±1.5 um (range 72 to 
77 um, n=20). Body oval (Fig. 1A), with posterior spinular row 
on ventral surface and on either side of caudal setae. Ventral 
surface with 2 paired patches of spinules posterior to labrum. 
Oval area of integument located posteriorly on ventral surface. 
Caudal rami represented by pair of naked setae. 

Antennule 3-segmented (Fig. 2A). First segment with 1 
spinulose seta anteriorly and group of spinules along outer 
margin. Second segment with 2 setae, distal one spinulose and 
slightly shorter than proximal plumose one; ornamented with 2 
spinular rows along ventral side. Third segment with 2 naked 
setae distally and 2 spinular rows along ventral margin. 

Antenna biramous (Fig. 3 A), with 2-segmented protopod 
comprising coxa and basis. Coxal gnathobase represented by 
large spine armed with spinular row distally. Basis with 3 small 
setae along inner margin, 2 of them closely set together in 
proximal third. Exopod 4-segmented; segment 1 large with 1 
spinulose seta and spinular row on outer margin; second 
segment with 1 long naked seta; third segment with 1 long naked 
seta and few tiny spinules on outer margin; fourth segment with 
naked seta proximally and 1 long spinulose seta plus 1 naked seta 
distally; outer margin with spinular row near proximal seta and 
single isolated setule apically. Endopod unsegmented; armed 
with 2 short inner setae and 2 long setae terminally. 

Mandible (Fig. 4A) biramous, with 2-segmented protopod 
comprising small coxa and large basis. Coxal gnathobase with 1 
naked seta. Inner margin of basis with 1 spinulose and 1 naked 
seta; outer margin with minute spinule. Exopod 4-segmented; 
first to third segments each with 1 plumose seta at inner distal 
angle; segments 3 and 4 with spinular row along outer margin; 
apex of fourth segment with long plumose, inner seta and short, 
naked, outer seta, the latter about as long as segment. Endopod 
2-segmented; first segment with two spinulose setae and spinular 
row on inner distal margin; second segment with 4 naked setae. 

Nauplius II 

Body length : mean ± standard deviation = 1 35 ± 7.84 um (range 
1 13 to 144 um, n=21), mean body width 88 ±4.5 um (range 83 to 
100 um, n=21). Body similar to first nauplius but larger and 
elongated caudally (Fig. IB); differing as follows: labrum (Fig. 



19C) with some spinules along lateral margin. Caudal region 
lacking spinular row adjacent to caudal setae. 

Antennule (Fig. 2B) armed with third seta on inner terminal 
edge of distal segment. Antenna (Fig. 3B) with additional small 
seta at base of large sword-shaped coxal gnathobasic seta. Basis 
with distal spinular row. Exopod 6-segmented, with 1 extra seta 
located on minute second segment (arrowed in Fig. 16A). 
Terminal segment of first nauplius subdivided; segment five with 
1 naked seta and additional spinular row distally. Endopod of 
antenna with 2 plumose setae of equal size and 1 shorter 
plumose seta terminally, plus 2 inner lateral naked setae. 

Coxa of mandible (Fig. 4B) with unilaterally spinulose seta. 
Additional naked seta present on inner margin of basis. First 
segment of endopod with one naked seta and one longer 
plumose seta but with no spinular row on inner distal margin. 
Second segment of endopod with 4 naked setae and 1 plumose 
seta. Base of first exopodal segment with additional naked seta. 

Maxillule appearing as strong plumose seta (arrowed in Fig. 
IB). 

Nauplius III 

Body length : mean ± standard deviation = 158 ± 6.08 um (range 
155 to 168 um, n=14), mean body width 105 ± 8.7 um (range 88 
to 117 um, n=14). Body (Fig. 1C) similar to second nauplius but 
larger, differing as follows: lateral spinular row on labrum 
consisting of much stronger spinules. Caudal margin with pair 
of strong plumose setae adjacent to longer naked setae. 
Posterior end of body trilobate. 

Third segment of antennule (Fig. 2C) acquiring extra naked 
seta on inner margin. Sixth exopodal segment of antenna (Fig. 
3C) with 1 naked inner seta, 1 strong spinulose subapical seta 
and 1 long spinulose apical seta. Mandible with minor changes 
in relative lengths of endopodal setae (Fig. 4C). 

Nauplius IV 

Body length : mean ± standard deviation = 181 ± 8.4 um (range 
166 to 192 um, n= 14), mean body width 1 25 ± 8.7 um (range 113 
to 144 um, n=14). Body (Fig. ID) pear-shaped, differing from 
Nauplius III as follows: caudal margin with pair of minute 
spines representing Anlagen of caudal setae. Posterior end of 
body trilobate and becoming elongated. 

Antennule (Fig. 2D) with a longer row of dentiform spinules 
distally on first segment. Third segment with 2 additional naked 
setae, plus extra row of long spinules midway along lateral 
margin. 

Coxa of antenna (Fig. 3D) with spinular row midway along 
outer margin. Basis with 4 setae along inner margin; outer 
margin with 2 spinular rows on anterior surface and 1 spinular 
row on posterior surface. Apical segment of exopod with 
spinular row distally. 

Basis of mandible with few mid-anterior surface spinules. 
Fourth segment of exopod with additional small seta distally 
(Fig. 4D). 

Maxillule (Fig. ID) bilobed with 2 spinulose and 1 naked seta 
on inner lobe. Outer lobe with 1 spinulose, 1 plumose and 1 long 
naked seta. 

Nauplius V 

Body length : mean ± standard deviation = 210 ± 8.49 um (range 
194 to 226 um, n=14), mean body width 138 ± 12.08 um (range 
122 to 157 um, n=14). Body (Fig IE) similar to nauplius IV, 
differing as follows: third pair of caudal setae larger. 



LIFE CYCLE OF PARACYCLOPS 



43 




Fig. 1 Pfimbriatus. Naupliar stages, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V; F, Nauplius VI. 
Antennules, antennae and mandibles omitted. Scale bar in |im. 



44 



S. KARAYTUG AND G.A. BOXSHALL 




Fig. 2 P. fimbriate. Naupliar antennules, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V- F Naupli 
VI. Scale bar in nm. 



LIFE CYCLE OF PARACYCLOPS 



45 




Fig. 3 P.fimbrialus. Naupliar antennae, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V; F, Nauplius VI. 
Scale far in |am. 



46 



S. KARAYTUG AND G.A. BOXSHALL 




Fig. 4 P. fimbriates. Naupliar mandibles, ventral view. A, Nauplius I; B, Nauplius II; C, Nauplius III; D, Nauplius IV; E, Nauplius V; F, Naupli 
VI. Scale bar in |im. 



LIFE CYCLE OF PARACYCLOPS 



47 



Hi 0* 




Fig. 5 P.fimbriatus. Dorsal view of Copepodid stages I-V. A, Female copepodid I; B, Female copepodid II; C, Female copepodid III; D, Male cope- 
podid IV; E, Female copepodid IV; F, Female copepodid V; G, Male copepodid V Scale bar in nm. 



48 



S. KARAYTUG AND G. A. BOXSH ALL 




Fig. 6 P.fimbriatus. Urosome of female copepodids, dorsal. A, Copepodid I; B, Copepodid II; C, Copepodid III; D, Copepodid IV; E, Copepodid 
V. Scale bar in um. 



First segment of antennule (Fig. 2E) with spinular row on 
outer margin. Third segment acquiring 3 small naked setae. 
Endopod of antenna (Fig. 3E) with additional small naked seta 
distally. Basis of mandible (Fig. 4E) with 4 setae. Maxillule (Fig. 
IE) with 1 new naked seta on inner lobe. 

Nauplius VI 

Body length : mean ± standard deviation = 241 ± 8.49 um (range 
255 to 224 um, n= 1 6), mean body width 1 52 ± 7.28 (range 143 to 
162 um). Body (Fig. IF) similar to nauplius V, differing as 
follows: innermost pair of caudal setae longest, middle pair of 
setae plumose, shortest setae naked and laterally located. 

Third segment of antennule (Fig. 2F) with total of 13 setae, 
including 5 new setae, plus ornamentation of several long 
spinules. Endopod of antenna with 9 setae (Fig. 3F), with 1 extra 
seta in proximal group. Mandible (Fig. 4F) with minor changes 
in ornamentation of basal setae. 



Maxillule with 1 new seta on inner lobe, and with 3 spinules on 
outer margin. Anlagen of post-maxillulary limb pairs apparent 
on ventral surface of body (Fig. IF). 

Description of Copepodid Stages 

Copepodid I 

Body length : mean ± standard deviation = 382 ±11.5 um (range 
373 to404um, n=10), mean body width 146 ± 5.7 urn (range 134 
to 152 um, n=10). Body 5-segmented (Fig. 5 A), comprising 
cephalothorax and 4 postcephalothoracic trunk somites. Second 
trunk somite with 1 seta at posterolateral angles representing 
third swimming legs. Antennule, antenna, mandible, maxillule, 
maxilla, maxilliped and first and second swimming legs present. 
Caudal rami (Fig. 6A) as broad as long. Each ramus armed with 
6 setae, ornamented with spinular row on ventral surface, 
outermost seta plumose with spinular row at base, 1 naked seta 
located on dorsal surface posteriorly, 1 short naked seta close to 



LIFE CYCLE OF PARACYCLOPS 



49 




Fig. 7 P.fimbriatus. Development of female antennules. A, Copepodid I; B, Copepodid II; C, Copepodid III; D, Copepodid IV; E, Copepodid V. 
Scale bar in jim. 



50 



S. KARAYTUG AND G. A. BOXSHALL 




Fig. 8 P. fimbriates. Development of copepodid antennae. A, Copepodid I; B, Copepodid II; C, Copepodid III; D, Copepodid IV; E, Copepodid V; 
F, Copepodid VI. Scale bar in urn. [Setal numbering scheme of Boxshall & Evstigneeva (1994) is used.] 



LIFE CYCLE OF PARACYCLOPS 



51 




Fig. 9 P. fimbriates. Copepodid I. A, Mandible, posteroventral; B, Maxillule, ventral; C, Labrum, ventral; D, Maxilla, anterior; E, Maxilliped, pos- 
terior. Scale bar in |im. 



outermost seta, 2 inner plumose setae distally and 1 naked 
seta on dorsal outer margin. Anal operculum ornamented 
spinular row. 

Antennule 5-segmented (Fig. 7A); aesthetasc at antero-distal 
angle of apical segment sharing common base with adjacent 
seta, as in all subsequent stages, including adult. Setal formula 3, 
3, 3 + 1 aesthetasc, 3,7+1 aesthetasc. 

Antenna 5-segmented (Fig. 8A); coxa unarmed; basis with 
vestigial exopod bearing 2 naked setae, one apically and one 
halfway along inner margin. Basis armed with 2 inner angle 
setae. Endopod 3-segmented; setal formula 1, 4, 6. Labrum (Fig. 
9C) with strong denticles along mid posterior margin. 

Mandible (Fig. 9A) with well developed gnathobase bearing 
row of sharp blades medially, and 1 spinulose seta at inner distal 
angle. Vestigial mandibular palp with 2 long plumose setae and 1 



naked seta distally. Inner posterior margin of mandible with two 
spinular rows. 

Maxillule with strong praecoxa and reduced 2-segmented 
palp (Fig. 9B). Praecoxal endite armed with 4 setae articulating 
at base and 4 spines fused to segment. Proximal segment of palp 
derived from coxa and basis, bearing 2 naked and 1 spinulose 
inner margin setae, plus outer seta representing exopod. Distal 
segment of palp representing endopod, armed with 3 naked 
setae. 

Maxilla (Fig. 9D) consisting of praecoxa, coxa, basis and 
2-segmented endopod. Praecoxa with single indistinct endite 
armed with 2 plumose setae. Coxa with inner seta at midlength 
and distal endite bearing strong plumose seta and naked seta 
apically. Basis with 3 setae distally. Endopod with 2 setae on 
proximal segment and 3 setae terminally. 



52 



S. KARAYTUG AND G.A. BOXSHALL 






50 



Fig. 10 P. fimbriate. Leg 1 , anterior. A, Copepodid I; B, Copepodid II; C, Copepodid III; D, Copepodid IV; E, Copepodid V; F, Copepodid VI 
(adult). Scale bars in urn. 



LIFE CYCLE OF PARACYCLOPS 



53 




Fig. 1 1 P.fimbriatus. Leg 2, anterior. A, Copepodid I; B, Copepodid II; C, Copepodid III; D, Copepodid IV; E, Copepodid V; F, Copepodid VI 
(adult). Scale bar in urn. 



54 



S. KARAYTUG AND G.A. BOXSHALL 




Fig. 12 P.fimbriatus. Leg 3, anterior. A, Copepodid II; B, Copepodid III; C, Copepodid IV; D, Copepodid V; E, Copepodid VI (adult). Scale bar in 
urn. 



Maxilliped (Fig. 9E) distinctly 4-segmented, comprising 
syncoxa, basis and 2-segmented endopod. Syncoxa armed with 
2 spinulose setae. Basis with 2 setae. Endopod segment 1 with 1 
plumose seta; segment 2 with 3 setae. 

Swimming legs 1 (Fig. 10A) and 2 (Fig. 11 A) each with 
2-segmented protopod. Intercoxal sclerites unornamented. 



Coxa of leg 1 with spinular row posteriorly. Basis of both legs 
with outer angle seta ornamented by spinular row at base, inner 
basal margin with fine spinules. Both legs with 1 -segmented 
rami. Inner margin of exopod and outer margin of endopod of 
both legs with row of pinnules. Spine and seta formula as 
follows: 



LIFE CYCLE OF PARACYCLOPS 



55 



Coxa 


Basis 


Exopod 


Endopod 


Leg 1 0-0 
Leg 2 0-0 


1-0 
1-0 


IV,4 
IV,3 


1,1+1,3 
1,1+1,3 


Copepodid II 









Body length : mean ± standard deviation = 455 ± 17.2 um (range 
439 to 488 um, n= 1 0), mean body width 1 68 ± 9. 1 um (range 1 52 
to 178 um, n=10). Differing from Copepodid I as follows: body 
(Fig. 5B) 6-segmented, comprising cephalothorax and 5 
postcephalothoracic trunk somites. Third trunk somite with 1 
seta on outer margin representing fourth swimming legs. Last 
somite with posterior margin spinular rows ventrally and 
laterally (Fig. 6B). 

Antennule with 3 spinules on first segment. Setal formula; 5, 4, 
4 + 1 aesthetasc, 3,7+1 aesthetasc (Fig. 7B). 

Antenna 4-segmented (Fig. 8B); basis with spinular row near 
inner margin. Inner setae set close together at distal angle. 
Vestigial exopod represented by spinulose seta. First endopodal 
segment with 1 seta, second with 5 setae along distal margin and 
bearing small spinular row on outer margin. Third endopodal 
segment with 6 setae distally, ornamented with spinular row on 
outer distal margin. 

Coxa of leg 1 with spinulose seta at inner distal angle (Fig. 
10B). Basis with spinulose seta on inner distal margin 
ornamented with spinular rows at base of inner seta and 
posteriorly, between exopod and endopod. Intercoxal sclerite 
ornamented with setules distally. First segment of exopod with 
spinular row along outer margin. First and second segments of 
endopod with pinnules along lateral margins. 

Coxa of leg 2 (Fig. 11B) with naked seta on inner distal 
margin; ornamented with spinular rows proximally on outer 
margin of both anterior and posterior surfaces and along outer 
margin. First segment of exopod with spinular rows laterally 
and proximally on posterior surface. Exopodal and endopodal 
segments with pinnules along inner and outer margins 
respectively. 

Third leg (Fig. 12A) with 2-segmented protopod; coxa with 
spinular row proximally near outer margin. Basis with naked 
outer seta; ornamented with tiny spinules at base and hairs along 
inner margin. Intercoxal sclerite unornamented. Exopod and 
endopod 1 -segmented, bearing pinnules along inner and outer 
margins respectively. Spine and seta formula as follows: 



aesthetasc. Antenna (Fig. 8C) with spinular row on inner margin 
of second segment proximally. Second endopodal segment with 
6 setae. Labrum with long spinular rows along lateral margins 
distally. 

Exopod of legs 1 (Fig. 10C) and 2 (Fig. 1 1C) with new seta on 
inner margin of first segment. Intercoxal sclerite of leg 2 
ornamented with rows of setules on anterior surface. Second 
endopodal and exopodal segments each with new seta. Coxa of 
third leg (Fig. 12B) with new seta at inner angle and bearing 
spinular row on outer margin. Intercoxal sclerite ornamented. 
Exopod and endopod 2-segmented. Leg 4 (Fig. 13A) with 
2-segmented protopod; coxa with spinular row on outer margin. 
Basis with outer angle seta and bearing spinular row on laterally. 
Exopod and endopod 1 -segmented. Intercoxal sclerite 
unornamented. Leg 5 (Fig. 6C) represented by 2 setae. Spine and 
seta formula as follows: 



Coxa Basis Exopod 



Endopod 



Legl 


0-1 


1-1 


I-1;III,5 


0- 1 ; 1 , 1+1,4 


Leg 2 


0-1 


1-0 


I-1;IV,5 


0-l;l, 1+1,4 


Leg 3 


0-1 


1-0 


I-0;IH,4 


0-1,1,1+1,3 


Leg 4 


0-0 


1-0 


IV,3 


1,1+1,3 



Female copepodid IV 

Body length : mean ± standard deviation = 573 ±19.5 um (range 
547 to 608 um, n=10), mean body width 200 ± 14 um (range 178 
to 221 um, n=10). Differing from copepodid III as follows: body 
(Fig. 5E) 8-segmented with 4-segmented urosome (Fig. 6D); 
spinular rows present mid-dorsally along posterior margins of 
urosomites 2 and 3. 

Antennule 6-segmented (Fig. 7D) with entire fourth segment. 
Setal formula; 11, 4, 5, 4 + 1 aesthetasc, 3, 7 + 1 aesthetasc. 
Antenna with 2 short spinular rows dorsally on inner side of first 
segment. Second endopodal segment with 7 setae (Fig. 8D). 
Mandible with spinular row on ventral margin of coxa. 
Maxilliped with spinular rows midway along outer margin and 
on anterior surface near inner setae. 

Legs 1-4 with 2-segmented rami (Figs 10D, 11D, 12C, 13B). 
Coxa of leg 4 with naked seta at inner angle proximally and 
spinular row on outer margin. Spine and seta formula as follows: 



Coxa Basis Exopod 



Endopod 



Coxa Basis Exopod 



Endopod 



Leg 1 0-1 
Leg 2 0-1 
Leg 3 0-0 



1-1 
1-0 
1-0 



1-0; 111,5 
1-0; 111,4 
IV,3 



0-1; 1,1+1,4 
0-1; 1,1+1,3 
1,1+1,3 



Copepodid III 

Body length : mean ± standard deviation = 51 1 ± 22.8 um (range 
478 to 534 um, n= 10), mean body width 1 79 ± 5.3 um (range 1 73 
to 186 um, n=10). Differing from copepodid II as follows: body 
(Fig. 5C) 7-segmented, comprising 4-segmented prosome 
bearing swimming legs 1 to 4, and 3-segmented urosome (Fig. 
6C). 

Antennule 6-segmented (Fig. 7C): fourth segment partly 
divided. Setal formula: 6, 2, 5, 4 + 1 aesthetasc, 3, 7 + 1 



Legl 


0-1 


1-1 


I-1;III,5 


0-1; 1,1+ 1,4 


Leg 2 


0-1 


1-0 


1-1;IV,5 


0-l;l,I+l,5 


Leg 3 


0-1 


1-0 


I-1;IV,5 


0-1; 1,1+ 1,4 


Leg 4 


0-1 


1-0 


I-0;IV,5 


0-l;l,II,3 



Leg 5 with 3 setae; leg 6 represented by 2 simple setae at 
posterolateral angle of second urosomite (Fig. 6D) 

Male copepodid IV 

Body length, (range 565 to 643 um, n=2), body width, (range 195 
to 204 um, n=2). Differing from female Copepodid IV as 
follows: anterior part of body (Fig. 5D) not much broader than 
posterior compared with female; constriction between somites 
bearing 4th and 5th legs (Fig. 14A) not as distinct. 

Appendages as in female fourth copepodid except as follows: 
antennule 6-segmented (Fig. 15A) but segment 3 broader. 
Segments 3 and 4 with distinctive short spiniform setation 



56 



S. KARAYTUG AND G.A. BOXSHALL 




Fig. 13 P.fimbriatus. Leg 4, anterior. A, Copepodid III; B, Copepodid IV; C, Copepodid V; D, Copepodid VI (adult). Scale bar in urn. 



elements (arrowed in Fig. 16D). Setal formula: 1 1, 4, 5 + spine, 4 
+ spine + 1 aesthetasc, 3, 7 + 1 aesthetasc. 

Female copepodid V 

Body length : mean ± standard deviation = 726 ± 38.47 um 
(range 647 to 769 urn, n=10), mean body width 246 ± 12.4 urn 



(range 217 to 260 um, n=10). Differing from female Copepodid 
IV as follows: body 9-segmented (Fig. 5F) with 5-segmented 
urosome (Fig. 6E); anal somite with dense spinular row along 
posterior margin extending from ventral surface to dorsal 
margin, either side of anal slit. Caudal rami about 3 times as 
long as broad. 
Antennule 7-segmented (Fig. 7E) : setal formula; 18, 6, 5, 2 + 1 



LIFE CYCLE OF PARACYCLOPS 



57 




Fig. 14 Pfimbriatus. Male. A, Copepodid IV urosome, dorsal; B, Copepodid V, dorsal; C, adult urosome, ventral; D, adult urosome, dorsal; E, 
Adult leg 5, ventral; F, Adult leg 6, ventral. Scale bars in urn. 



58 



S. KARAYTUG AND G.A. BOXSHALL 




Fig. 15 P.fimbriatus. Male antennules, lateral. A, Copepodid IV; B, Copepodid V. Scale bar in \m\. 



LIFE CYCLE OF PARACYCLOPS 



59 




Fig. 16 Scanning electron micrographs of P. fimbriatus. A, Nauplius VI antenna, ventral; arrow indicates minute second exopodal segment; B, 
Adult female antennule, aesthetasc on segment 5; C, Male copepodid V antennule, dorsal; D, Male copepodid IV antennule, ventral; arrow indi- 
cates the spine on segment 4; E, Adult female antennule, double fusion on terminal segment, lateral. Scale bars A = 20 um, B = 5um, C = 40 urn, D 
= 25 um, E = 5 um. 



aesthetasc, 2, 3, 7 + 1 aesthetasc. Second endopodal segment of 
antenna (Fig. 8E) with 8 setae; distal seta VIII stronger than 
others. Praecoxal arthrite of maxillule with 5 setae. Proximal 
spinulose seta ornamented with long spinules. Praecoxa of 
maxilla with short spinular row on outer margin. Basis of 
maxilliped with 2 spinular rows laterally. 



Leg 1 (Fig. 10E) with 3-segmented exopod and endopod, 
bearing spinular row posteriorly on second segment of exopod. 
Intercoxal sclerite with anterior spinular rows distally. Leg 2 
(Fig. 1 IE) with praecoxa bearing spinular row on outer margin. 
Exopod and endopod 3-segmented: first segment of exopod 
with proximal spinular row anteriorly, second exopodal and 



60 



S. KARAYTUG AND G.A. BOXSHALL 





Fig. 17 P. fimbriatus. A, Adult female, dorsal; B, Adult female antennule, dorsal; C, Adult male, dorsal. Scale bars in urn. 



endopodal segments with spinular rows posteriorly. Leg 3 (Fig. 
12D) with 3-segmented exopod and endopod bearing posterior 
spinular rows on first and second segments of exopod and 
second segment of endopod. Intercoxal sclerite with transverse 
spinular row on anterior surface. Leg 4 (Fig. 1 3C) with praecoxa 
bearing spinular row on outer margin. Coxa with posterior 



spinular row near outer proximal angle, spinular row on 
midanterior surface and very fine spinules along posterior 
margin. Exopod and endopod 3-segmented: first and second 
segments of exopod with anterior spinular rows distally. 
Intercoxal sclerite with transverse spinular row on anterior 
surface. Spine and seta formula as follows: 



LIFE CYCLE OF PARACYCLOPS 



61 




Fig. 18 P. fimbriates. Adult female. A, urosome, ventral; B, urosome, dorsal. C, leg 5, ventral. Scale bars in urn. 



62 



S. KARAYTUG AND G.A. BOXSHALL 



Coxa Basis Exopod 



Endopod 



Legl 


0-1 


1-1 


I-1;I-1;III,1,4 


0-1 ;0-l; 1,1,4 


Leg 2 


0-1 


1-0 


I-1;I-1;III,I,5 


0-l;0-2;l,I,4 


Leg 3 


0-1 


1-0 


I 1;I-1;III,I,5 


0-1 ;0-2; 1,1,4 


Leg 4 


0-1 


1-0 


I-1;I-1;III,1,4 


0-1 ;0-2; 1,11,2 



Leg 6 represented by 1 plumose and 1 naked seta at 
posterolateral angle of second urosomite (Fig. 6E) 

Male Copepodid V 

Mean body length: mean ± standard deviation = 695 ± 44.83 urn 
(range 634 to 743 um, n=7), mean body width 216 ± 8.1 urn 
(range 200 to 221 .73 um). Differing from male Copepodid IV as 
follows: body 9-segmented: first to fourth urosomites 
ornamented along posterior margins (Fig. 14B). Anal somite 
densely furnished with spinules along distal margin and 
extending either side of anal operculum. 

Appendages as in fourth copepodid except as follows: 
antennule (Fig. 15B) 7-segmented; proximal 3 segments much 
enlarged, swollen (Figs 16C) first segment with one modified 
spiniform seta (stippled on Fig. 15B); segment 3 with 6 similarly 
modified elements and segment 4 with 1 such element. Setal 
formula; 19+1 spine, 5, 5 + 6 spines, 2 + 1 aesthetasc + 1 spine, 2, 
3,7+1 aesthetasc. 

Leg 6 (Fig. 14B) represented by 3 setae, not visible in dorsal 
view. 

Adult female 

Body length: mean ± standard deviation = 935 ± 61.07 um (range 
833 to 1013 um, n=10), mean body width 307 ± 14.9 um (range 
291 to 330 um, n=10). Body (Fig. 17A) comprising 4-segmented 
prosome and 5-segmented urosome. Prosome with 
cephalothorax and 3 free pedigerous somites decreasing in width 
from anterior to posterior. Cephalothorax narrowing anteriorly; 
widest aproximately in middle. Third and fourth pedigerous 
somites with lateral groups of bristles at posterolateral angles. 
Urosome (Figs 18A,B) consisting of 5th pedigerous somite, 
genital double-somite and 3 free abdominal somites. Genital 
double-somite about as long as broad. Genital double-somite 
and first 2 free abdominal somites with surface ornamentation 
dorsally and ventrally : fifth pedigerous somite with 
ornamentation along posterior margin. Anal somite with 
spinular row ventrally extending round to anal operculum (Fig. 
19A). Anal operculum smooth; row of spinules present in anal 
cleft either side of midline (Fig. 19D). 

Caudal rami about 3.5 times longer than broad with 
ornamentation comprising rows of pits or cuticular depressions 
on ventral surface (Fig. 19B). Generally held wide apart, slightly 
divergent. Caudal rami with 6 setae; seta I missing; outer lateral 
seta (III) with spinular row at base extending dorsally and 
ventrally; small seta (II) on dorsolateral surface with spinular 
row behind it and extending ventrally. 

Antennule 8-segmented (Fig. 17B), quite short. First segment 
with spinular row ventrally. Segment 2 with partial suture line. 
Fourth segment longest. Segment 5 distinctive with short 
aesthetasc (Fig. 16B); apical segment with aesthetasc fused to 
adjacent seta at base (Fig. 16E). Setal formula: 8, 12, 6, 5, 2 + 1 
aesthetasc, 2, 3, 7 + 1 aesthetasc. One element on seventh 
segment possibly a setiform aesthetasc (see discussion below). 

Antenna 4-segmented (Fig. 8F), comprising coxobasis and 
3-segmented endopod. Coxobasis with complex ornamentation 



as figured and armed with 2 inner setae, and 1 outer spinulose 
seta representing exopod. First endopodal segment with inner 
distal seta and midsurface spinular row. Second endopodal 
segment with 9 setae, of which 5 on inner margin and 4 arranged 
along inner part of distal margin; segment ornamented with 
spinules along outer margin. Third endopodal segment armed 
with 7 setae around apex; segment ornamented with spinular 
row along outer margin. 

Labrum with complex ornamentation on ventral surface (Fig. 
20A). Anterior part broader than posterior part. Posterior 
margin forming strong teeth. 

Mandible (Fig. 20B) consisting of well developed coxal 
gnathobase and reduced palp. Gnathobasic blades mostly 
simple, dorsal seta with spinules along inner rim. Palp 
represented by 3 setae, one long and spinulose, the other 
plumose. Third seta short and naked. Central surface of coxa 
with 3 spinular rows and another spinular row on margin at base 
of gnathobase. 

Maxillule (Fig. 20C) consisting of powerful praecoxa and 
reduced 2-segmented palp. Praecoxal arthrite armed with 7 setae 
articulating at base and 4 spines fused to segment; proximalmost 
seta spinulose, spines naked. Proximal segment of palp derived 
from coxa and basis, bearing 2 naked and 1 spinulose inner 
margin setae, plus outer seta representing exopod. Distal 
segment of palp, representing endopod, armed with 2 setae with 
spinules along margin and 1 naked seta. 

Maxilla 5-segmented (Fig. 20D), comprising praecoxa, coxa, 
basis and 2-segmented endopod. Praecoxa with spinular row on 
outer margin. Praecoxal endite with 2 setae, 1 of which 
spinulose. Coxa with proximal endite represented by single seta, 
distal endite with well developed process carrying strong 
spinulose seta and naked seta apically. Basis drawn out into 
powerful spinulate claw and armed with strong accessory claw 
with spinular row along convex margin and naked seta. First 
endopodal segment carrying 2 spinulose setae, second carrying 3 
setae. 

Maxilliped 4-segmented (Fig. 20E), comprising syncoxa, 
basis and 2-segmented endopod. Syncoxa armed with 3 inner 
margin setae representing endites; ornamented with spinular 
row near middle of inner margin and 2 spinules on outer margin. 
Basis armed with 2 inner setae, 1 of which with spinules; 
ornamented with 2 transverse rows of spinules near outer distal 
angle and another spinular row midway along outer margin. 
First endopodal segment bearing claw-like seta with 5 spinules at 
midlength. Second endopodal segment with 3 setae, 2 of which 
bearing spinules midway; outermost naked. 

Legs 1 to 4 each with complex ornamentation on anterior 
and posterior surfaces of coxa as figured; also ornamented on 
intercoxal sclerite. Legs 1 to 3 with spinular rows on posterior 
surface of exopodal segments 1 and 2, and endopodal segment 
2. 

Leg 1 (Fig. 10F) with 3-segmented protopod. Praecoxa 
represented by triangular sclerite at outer proximal angle. Coxa 
with inner plumose seta. Basis with outer angle seta and setiform 
spine on inner margin; bearing spinular row on posterior 
surface. Both seta and spine with spinular rows at bases. 

Legs 2 (Fig. 1 IF) and 3 (Fig. 12E) with 3-segmented 
protopods. Coxa with inner setiform spine. Outer seta on basis 
with spinular row at base. First segment of exopod with spinular 
row on anterior surface. 

Leg 4 (Fig. 13D) with 3-segmented protopod. Praecoxa 
represented by thin, hooped sclerite. Coxa with inner setiform 
spine. Basis with outer seta bearing spinular row at base; 
spinular row on inner margin anteriorly. Segments 1 and 2 of 



LIFE CYCLE OF PARACYCLOPS 




Fig. 19 Scanning electron micrographs of P. fimbriatus. A, Adult female, anal operculum, lateral; B, adult female, caudal rami showing ornamenta- 
tion of pits in integument; C, Nauplius II, ventral; D, Adult female, anal operculum, dorsal. Scale bars A = 20 um, B = 40 um, C = 60 um, D = 25 
Jim. 



exopod with spinular row on distal margin anteriorly. Spine and 
seta formula as follows: 



Coxa Basis 



Exopod 



Endopod 



Leg 1 0-1 

Leg 2 0-1 

Leg 3 0-1 

Leg 4 0-1 



1-1 

1-0 
1-0 
1-0 



I-1;I-1;III,1,4 
I-1;I-1;III,I,5 
I-1;I-1;III,I,5 
I-1;I-1;II,I,5 



0-l;0-l;l,I,4 
0-l;0-2;l,I,4 
0-1 ;0-2; 1,1,4 
0-1 ;0-2;l, 11,2 



Leg 5 (Fig. 18C) comprising single free segment, armed with 1 
well developed outer spinulose seta, 1 strong inner spine 
ornamented with spinules, and 1 plumose seta in middle. Leg 6 
(Fig. 18 A) represented by 1 naked seta and 1 tiny spinule 
dorsolateral^. 

Adult male 

Body length: Mean ± standard deviation = 732 ± 47.45 um 
(range 673 to 769 um, n=10), mean body width 237 ± 7.6 um 



S. K.ARAYTUG AND G.A. BOXSH ALL 




Fig. 20 P. fimbriatus. Adult female. A, Labrum, ventral; B, Mandible, posteroventral; C, Maxillule, ventral; D, Maxilla, posterior; E, Maxilliped, 
anterior. Scale bar in urn. 




Fig. 21 P. fimbriatus. Male antennule. A, dorsal view. B, Detail of segments XI to XVI, anterior; C, Detail of segments XIII to XX, lateral view. 
Scale bar in nm. 



66 



S. KARAYTUG AND G.A. BOXSHALL 




Fig. 22 Scanning electron micrographs of P.fimbriatus. Adult male antennule. A, Dorsal view, with segment 5 arrowed; B, Lateral view, with sheath 
on segment XV arrowed; C, Modified seta on proximal segment; D, Segment proximal to geniculation, with pore on modified element arrowed. 
Scale bars A = 30 urn, B = 25 urn, C = 6 urn, D = 5 urn. 



(range 226 to 247 um, n=10). Body (Fig. 17C) differing from 
adult female as follows: urosome 6-segmented, comprising fifth 
pedigerous, genital and 4 free abdominal somites; 
ornamentation of spinule rows as in Fig. 14D. Genital somite 
bearing paired genital apertures ventrally (Fig. 14C). 

Caudal rami about 2.5 times longer than broad, ventral 



surface unornamented. Large medial seta (V) relatively longer 
than in female. 

Antennule 15-segmented (Figs 21A-C). Segment 1 armed 
with 9 setae; 1 seta large and modified (Fig. 22C), by 
ornamentation of strong spinules in proximal and mid sections, 
tapering to fine point distally; segment ornamented with curved 



LIFE CYCLE OF PARACYCLOPS 



67 



row of long spinules and row of minute spinules. Segment 2 with 
4 setae. Segment 3 with 2 setae. Segments 4 and 5 partly fused, 
with segment 5 (arrowed in Fig. 22A) defined only on dorsal side, 
not ventrally. Segment 4 with 2 setae; segment 5 with 2 setae. 
Segment 6 with 2 setae. Segments 7, 8 and 9 separated from each 
other by extensive arthrodial membrane: segment 7 with 2 setae, 
segment 8 with 2 setae, and segment 9 with 2 setae plus 
aesthetasc. Segment 10 (= ancestral segment XV) produced on 
one side into extensive sheath (arrowed in Fig. 22B) enclosing 
segment 1 1 ventrally: armed with 2 setae, 1 ornamented with 
long spinules unilaterally. Segment 1 1 bearing curved seta 
ornamented with double row of strong denticles, plus 1 naked 
seta. Segment 12 partly fused to segment 13; armed with short 
seta ornamented with 2 rows of fine spinules, plus short naked 
seta. Segment 13 partly subdivided by partial suture: armed with 
short spinulate seta proximally, 2 short naked setae, plus 1 
modified element attached to segment by short stalk, main part 
of element lying along surface of segment and ornamented with 
longitudinal ridges and small central pore (arrowed in Fig. 22D). 
Geniculation located between segments 13 and 14. Segments 14 
and 15 partly fused, forming curved subchela-like section: 
segment 14 armed with 1 seta, 1 aesthetasc and 2 modified 
elements each ornamented with longitudinal ridges and a central 
pore, as distal element on segment 13. Apical segment tapering 
distally; armed with 1 1 setae and 1 aesthetasc, mostly 
originating on outer (= posterior) surface. 

All other appendages as in female except for fifth (Fig. 14E) 
and sixth legs (Fig. 14F). Outer spinulose seta of leg 5 
ornamented with some long setules distally. Sixth legs forming 
opercular plates bearing row of large spinules along ventral 
surface; armed with 1 inner spine, 1 well developed spinulose 
seta and 1 inner naked seta. 



DISCUSSION 

The number of naupliar instars in the Cyclopoida has been the 
subject of some controversy but it is clear, as Elgmork & 
Langeland (1970) strongly indicated, that there are normally 6 
naupliar instars. This is supported by recent works on free-living 
freshwater Cyclopoida by Dahms & Fernando (1992, 1993, 
1994) and by our data. The most difficult distinction is between 
nauplius IV and V and these stages have often been confused. 

The complete naupliar sequence of P. fimbriatus was 
previously described by Ewers (1930) and Dukina (1956). 
Nauplius I and II were also described by Gurney (1933) but none 
of these provides setation counts of sufficient accuracy. Ewers 
(1930) described 6 stages but our descriptions differ as follows: 
antennule is 3-segmented not 4-segmented; antennary exopod is 
4-segmented at N I and becomes 6-segmented, rather than 
remaining 4-segmented; caudal rami of N IV are represented by 
2 pairs of setae and 1 pair of minute seta, rather than just 2 pairs 
of setae; ventral body surface is ornamented with spinular rows 
throughout the nauplius phase. In general, however, Ewers' 
drawings are so small that it is not worthwhile making detailed 
comparisons of appendage setation patterns. 

Six naupliar stages were also described by Dukina (1956). Our 
descriptions differ from Dukina's as follows: antennule is 
3-segmented; antennary exopod is 4-segmented at N I (given as 
5-segmented by Dukina). Dukina's descriptions of appendage 
setation also lack sufficient detail for meaningful comparisons. 

Gurney (1933) described the first 2 naupliar stages, but our 



findings indicate that the caudal rami of N II are represented by 
a pair of setae not by 2 pairs as illustrated by Gurney. Apart from 
this discrepancy our results differ only in that the exopod of 
antenna is described as 3-segmented rather than 4-segmented as 
in our material. 

The copepodid stages of P. fimbriatus were also partly 
described by Gurney (1933). Although our results are in 
substantial agreement, for example, with antennulary 
segmentation throughout the copepodid phase, Gurney's 
drawings are not sufficiently accurate to permit comparisons of 
segmental setation. 

Analysis of the antennulary setation patterns of adult male 
Paracyclops permits the identification of the pattern of segmental 
homologies. The basic armature of each antennulary segment in 
copepods is 2 setae plus one aesthetasc (Giesbrecht, 1 892), with a 
few exceptions as identified by Huys & Boxshall ( 1 99 1 ). Using this 
basic pattern, the 9 setae on the first segment of male P.fimbriatus 
indicate that it can be identified as representing 5 ancestral 
segments (segments I-V). The second segment, with 4 setae, can 
similarly be identified as derived from 2 ancestral segments 
(VI-VII). The third to twelfth segments all represent single 
ancestral segments (VIII to XVII), as indicated by the presence of 
a maximum of 2 setae on each. The fourth and fifth are 
incompletely separated (Fig. 22A) but we have treated them as 
distinct. The thirteenth segment has only 4 setae but is here 
identified as representing 3 ancestral segments (XVIII-XX). This 
decision is based on the presence of the neocopepodan 
geniculation between the thirteenth and fourteenth segments 
which unequivocally identifies the segmental boundary involved as 
XX to XXI, and on comparison with other cyclopids such as 
Etayte robusta Giesbrecht, 1900. In E. robusta males segment 
XVIII is separate and carries a long naked seta and a short 
spinulose seta, segment XIX-XX carries a short spinulose seta and 
a modified spine proximally and a slender seta distally (Huys & 
Boxshall, 1991). In P. fimbriatus the proximal part of the triple 
segment is defined by a partial suture marking the original plane 
between segments XVIII and XIX-XX. This part carries only a 
single spinulose seta and lacks the long seta; the distal part 
representing XIX-XX carries the same setation as in E. robusta. 
This confirms our interpretation of the thirteenth segment as a 
triple segment (XVIII-XX). The fourteenth and fifteenth 
segments, lying distal to the geniculation represent ancestral 
segments XXI-XXIII and XXIV-XXVIII, exactly as Huys & 
Boxshall ( 1 99 1 ) found for E. robusta. 

Compound antennulary segments, such as the first and 
second segments of P. fimbriatus, were simply referred to as 
'fused' by Huys and Boxshall (1991) in their comparative 
analysis of antennulary segmentation patterns in all copepod 
orders, although such compound segments could be the result of 
two different developmental processes: 

1) secondary fusion of segments that were separated earlier 
during ontogeny 

2) failure of separation during development. 

The compound first and second segments of the male 
antennule of P. fimbriatus result from the second process, the 
failure to separate. In contrast, the compound apical segment 
results from the secondary fusion of the sixth, seventh and 
eighth segments of the copepodid V stage. (The eighth segment 
of the copepodid V was already a compound segment, 
representing three ancestral segments XXVI-XXVIII which are 
not separately expressed by any known member of the order 
Cyclopoida). 

Vertical tracking of the segmental boundaries as identified by 
their setation elements allows us to identify the homologies of 



68 



S. KARAYTUG AND G.A. BOXSHALL 



o 



\) 



E „ 



o 



o 



o 



> 

o 



> 

o 



> 

o 



> 

o 



> 



> 
o 



-. 


■a 




Ctl 






'.1 


II 


■i 




CO 
OS 


- 



CD 












03 




CC 




05 


+ 




-»- 




+ 


h- 




h- 




h- 
















03 




03 




03 


+ 




+ 




+ 


r^ 




i^ 




r^ 



CO 




co 




CO 



CO 




CO 




co 








CD 




CD 


03 




CO 




CO 


+ 




+ 




+ 


CO 




^t 




^t 



CO 




^J- 




CD 




CM 




CM 










CO 




CD 




CD 


+ 




CO 




CO 


^" 




+ 
CM 




+ 
CM 



ID 




LO 




id 



^t- 




CD 




CD 



> 

X 
X 

J_ 

> 

X 
X 



> 

X 
X 



X 
X 

_!_ 

X 
X 



X 
X 



> 

X 



> 

X 

I 



CO 



if) 



CD 



1— 




CO 

1- 












CO 




03 


-h 




+ 


r^ 




r^ 



CO 




CO 




CM 



CD 


Cl> 


co 


C 


+ 


a. 


CM 


CO 



O) 







V) 






<D 






c 






a 






co 


CD 




CD 

+ 
ID 




CM 



CM 



CM 




CM + 



0)1 > 
«J X 



CM 



™J X 



CM £ 







(M 
CM 


X 
1 
X 


CO 

c 








a 

CO 




CM 


> 










-1- 




^ 


> 
> 




-C 


o 


-, 


BQ 


o 


3 


CO 


CO 


■a 


3 


cd 


X 


c 


>» 


c 


£3 








-3 
O 


E 


n 


f,; 




SI) 


p. 


Cm 


5) 


O 


= 


l: 


r 





a 




<i> 


rrt 




:) 


■- 





n 






00 

c 




* 


09 



en 


= 

Of 


lo 
u 

X 
1) 
en 

C 


■a 

i> 

c 
E 




r-n 





u 


J2 




•— 


00 





n 


CO 


"5 


u, 


'-J 


~. 






'.) 


CO 


-C 


^3 





T3 


P. 


O 


>-■ 


a. j= 


u 


E 


O 


o 


tj 


£ 










ed 


c 


e 


o 










CO 


T| 


'5b 







u 


o 


-a 


6 
o 


3 

u 




't} 


-1 


c 






C 








fi 




bfl 


I) 


u 


fc 


fc 


3 


ed 


C 


=) 


C3 
E 


c 


o 


tu 


a: 


c 


1 ' 


m 


Crt 


o 


2? 


a 




to 






'— 


B) 


a 


c 


p. 




E 


11) 


o 


r-1 


o 




u 


CO 






a 


nt 


e 


i) 


u 


-= 


-= 




Vi 




to 


■O 




c 


1 


CO 

g 


CO 


a; 


1)' 




S3 






n 


OJ 


<N 


111 


sr 


O 






u. 





LIFE CYCLE OF PARACYCLOPS 



69 



the segments of the female antennule and of the antennules 
throughout the copepodid phase of development. These results 
are presented in schematic form in Figure 23 which visually 
indicates homologous segmental boundaries expressed in 
successive copepodid stages. The aesthetasc derived from 
ancestral segment XXI provides a reference point throughout 
the copepodid phase: from the third segment of the first 
copepodid, to the fifth segment of the adult female and the 
fourteenth segment of the adult male. This, in concert with other 
setation features, confirms the unequivocal identification of the 
XX to XXI articulation (marked by the large arrow) in all stages. 
Identification of the homology of the more proximal segments 
relies on the presence of 1 1 setation elements on the third 
segment of the male copepodid V stage. These 1 1 elements 
testify to the segments derivation from 6 unseparated ancestral 
segments (XV-XX) and indicates that the boundary between the 
second and third segments represents the XIV to XV 
articulation. Similarly, the presence of 5 setae on the second 
segment indicates that the boundary between first and second 
segments represents the XI to XII articulation. The remaining 
segmental homologies in all stages, including the adult female, 
can now be identified by tracking vertically through the male 
stages back to copepodids I and II and then forwards again to 
the adult female. 

The seventh segment of the female antennule is identified as 
homologous with a single ancestral segment (XXV). The 
presence of 3 setae on this segment therefore requires 
explanation since the basic armature of segment XXV is 1 
posterior margin seta plus 1 seta and 1 aesthetasc on the anterior 
margin (Huys & Boxshall, 1991). We suggest that one of the 
anterior elements may be a setiform aesthetasc and this should 
be investigated ultrastructurally. 

This is the first time that the homologies of all antennulary 
segments have been identified in all copepodid stages of both 
sexes of any cyclopoid copepod, with reference to the 
hypothetical 28-segmented antennule of the ancestral copepod 
(Huys & Boxshall, 1991). Within the family Cyclopidae 
antennulary segmental numbers are frequently reduced and, 
indeed, many genera are characterised by their low segment 
numbers. The possible involvement of heterochrony in the 
reduction of antennulary segmental numbers has already been 
noted. Gurney (1933) suggested that 'the reduced numbers of 
antennulary segments in some species may be accounted for as 
the persistence of a larval character in the adult', pointing out 
that during development the last copepodid (Co V) typically has 
an 11 -segmented antennule and that this number is commonly 
found in adults, as for example in Microcyclops. 

Several authors working on cyclopids were aware of the 
importance of determining homologies between the different 
generic segmentation patterns and made detailed comparisons. 
Gurney (1933) reviewed earlier work by Claus (1893), Manfredi 
(1923), Lucks (1929) and Gelmini (1928) on the sequence of 
segmental subdivision during development in Cyclops species. He 
presented a tabular system showing derivations of a 1 7-segmented 
antennule and confirmed this pattern using his own data on C. 
strenuus. The strict determination of ancestral homologies of 
antennulary segments provides new insight into the rather 
confused world of cyclopid systematics at the generic level. 

The scheme of segmental development (Fig. 23) indicates that 
at the moult from copepodid II to copepodid III it is the second 
segment that subdivides. This contrasts with the scheme 
presented for P. fimbriatus by Gurney (1933) who showed the 
first segment subdividing at this moult. We base our 
interpretation on the positions and relative lengths of the setae 



on these segments. In particular a relatively long seta is 
positioned at the anterodistal angle of the first segment in 
copepodid II and 2 similarly long setae are present at the same 
position in copepodid III. Since no other setae of this length are 
present we interpret this as evidence of the constancy of the 
boundary marked by this seta and therefore conclude that the 
second segment has subdivided. 

The setation of the second endopodal segment of the antenna 
increases progressively through the copepodid stages of P. 
fimbriatus in a very regular manner. The first copepodid stage 
possesses 4 setae distributed around the inner-distal angle of the 
segment (Fig. 8A). The seta which is located on the angle is slightly 
stouter than the other 3 and is here identified as seta VIII, using the 
numbering scheme proposed for another member of the family 
Cyclopidae by Boxshall & Evstigneeva (1994). The more distal seta 
is then identified as seta IX and the 2 more proximally located setae 
on the inner margin, as setae VII and VI. These 4 setae are 
presumably homologous with the proximal group of 4 setae 
present on the margin of the unsegmented endopod of the sixth 
nauplius stage (Fig. 3F). At each successive moult through the 
copepodid phase one additional seta is added proximally on the 
inner margin of the segment (Figs 8A-F). Thus at copepodid II a 
fifth seta (seta V) is added, at copepodid III a sixth seta (seta IV), 
and so on, until the final moult to adult (= copepodid VI) at which 
the ninth and final seta (seta I) is added. 

The progressive development of the setation suggests that 
reductions in numbers of setae on the second endopodal segment, 
which are common within the family Cyclopidae, may be 
interpreted as resulting from heterochronic events. Cryptocyclops 
bicolor (Sars, 1 863), for example, is a cyclopid with only 7 setae on 
the second endopodal segment. This number is typical of the 
copepodid IV stage in both Paracyclops and the presumed 
ancestral cyclopid stock, and may be interpreted as evidence that a 
neotenic event within the Cryptocyclops lineage has interrupted the 
progressive addition of setae. 

The ontogeny of copepodid stages and examination the 
patterns of leg formation offer some evidence for inferring 
copepod phylogenetic relationships (Ferrari, 1988). A common 
pattern of development for legs 1-4, exhibited by at least 20 
genera, was recognized by Ferrari. The pattern of development 
for the swimming legs of P. fimbriatus is in accordance with this 
common pattern as follows: 



Legs 


1 


2 


3 


4 


N 


Tb 


1~B 






I 


i + i 


1 + 1 


Tb 




II 


2+2 


2+2 


1 + 1; 


1 B 


III 


2+2 


2+2 


2+2; 


1 + 1; 


IV 


2+2 


2+2 


2+2; 


2+2; 


V 


3+3 


3+3 


3+3; 


3+3; 


VI 


3+3 


3+3 


3+3; 


3+3; 



(Where N = nauplius; Roman numerals = copepodid stages; PB = 
primary leg bud; 1+1 = reorganized leg with 1 -segmented exopod and 
endopod; 2+2 = leg with 2-segmented exopod and endopod; 3+3 = leg 
with 3-segmented exopod and endopod). 



Acknowledgements. We are grateful to Dr Rony Huys for his helpful 
comments on drawing techniques and on the manuscript. S. Karaytug 
would also like to thank Dr Steve Alston for his assistance in the 
laboratory. This research has been supported by a postgraduate grant 
from the University of Balikesir, Turkey to S. Karaytug. 



70 



S. KARAYTUG AND G.A. BOXSHALL 



REFERENCES 



Boxshall, G.A. & Evstigneeva, T. 1994. The evolution of species flocks of copepods 

in Lake Baikal: a preliminary analysis. Archiv fur Hydrobiologie, Beiheft 

Ergebnisse der Limnologie 44: 235-245. 
Claus, C. 1893. Die Antennen der Pontelliden und das Gestaltungsgesetz der 

mannlichen Greifantennen. Sitzungsberichte der Akademie der Wissenschuften. 

Mathematisch-Naturwissenschaftliche Classe, Wien 101: 848-866. 
Dahms, H.-U. 1991a. Usefulness of postembryonic characters for phylogenetic 

reconstruction in Harpacticoida (Crustacea, Copepoda). Proceedings of the 

Fourth International Conference on Copepoda. Bulletin of Plankton Society of 

Japan, Special Volume: 87-104. 
Dahms, H.-U. 1991b. Naupliar development of Harpacticoida (Crustacea, 

Copepoda) and its significance for phylogenetic systematics. Microfauna Marina 

6: 169-272. 
Dahms, H.-U. & Fernando, C.H. 1992. Naupliar development of Mesocyclops 

aequatorialis similis and Thermocyclops consimilis (Copepoda: Cyclopoida) 

from Lake Awasa, a tropical rift valley lake in Ethiopia. Canadian Journal of 

Zoology 70: 2283-2297. 
Dahms, H.-U. & Fernando, C.H. 1993. Naupliar development of Mesocyclops cf 

thermocyclopoides Harada, 1931 and Thermocyclops decipiens (Kiefer, 1929) 

(Copepoda: Cyclopoida) from Beira Lake, Sri Lanka. Journal of Plankton 

Research 15: 9-26. 
Dahms, H.-U. & Fernando, C.H. 1994. Redescription of female Macrocyclops 

fuscus (Jurine, 1820) (Copepoda, Cyclopoida) from Ontario with a description 

of naupliar stages. Journal of Plankton Research 16: 9-2 1 . 
Dukina, V.V. 1956. Specific differences in the larvae of Cyclopidae. Zoologicheskii 

Zhurnal 35: 680-690. 
Dussart, B.H. 1969. Les Copepodes des eaux continentales d Europe Occidentale. II. 

Cyclopoides el Biologic. N. Boubee & Cie, Paris. 290pp. 
Dussart, B. H. & Defaye, D. 1985. Repertoire mondial des Copepodes Cyclopoides. 

Centre National de la Recherche Scientifique. 236pp. 
Einsle, U. 1971 . Copepodes Libres. Free-living Copepods. Exploration 

hydrobiologique du Bassin du lac Bangwcolo el du Luapula 1 3: 1 -74. 
Elgmork, K. & Langeland, A.L. 1970. The number of naupliar instars in 

Cyclopoida (Copepoda). Crustaceana 18: 277-288. 
Ewers, L.A. 1930. The larval development of freshwater Copepoda. The Ohio 

State University Press, Columbus. Contributions. Franz Theodore Stone Institute 

of Hydrobiology 3: 1-43. 
Ferrari, F.D. 1988. Developmental patterns in numbers of ramal segments of 

copepod post-maxillipedal legs. Crustaceana 54: 256-293. 
Fischer, S. 1853. Beitrage zur Kenntnis der in der Umgregend von St Petersburg 



sich findenden Cyklopiden (Forsetzung). Bulletin de la Societe Imperial des 

Naturalistes de Moscou 26: 74-100. 
Gelmini, G. 1928. Contributo alia conoscenza dello sviluppo larvale di Cyclops 

leuekarti, Claus. Natura, Milano 19: 89-96. 
Giesbrecht, W. 1892. Systematik und Faunistik der pelagischen Copepoden des 

Golfes von Neapel und der angrenzenden Meeres-Abschnitte. Fauna und Flora 

des Golfes von Neapel 19: 1-83 1 . 
Gurney, R. 1933. British Freshwater Copepoda. Vol. III. The Ray Society, London, 

384pp. 
Huys, R. & Boxshall, G.A. 1991. Copepod Evolution. The Ray Society. London. 

468pp. 
Ito, T. 1954. Cyclopoida Copepods of Japanese subterranean waters. Report of the 

Faculty of Fisheries Prefeetural University of Mie 3: Y12-A 16. 
Kiefer, F. 1929. Zur Kenntnis einiger Artengruppen der Susswasser-Cyclopiden. 

Zeitschrift fur Wissenschaftliche Zoologie 133: 1-56. 
Lindberg, K. 1941. Cyclopides (Crustaces Copepodes) de l'lnde. VIII. Cyclops 

Muller; IX. Paracyclops Claus & Ectocyclops Mullen X. Acanlhocyclops Kiefer 

& Diacyclops Kiefer. Records of the Indian Museum. Calcutta 43: 471-496. 
Loffler, H. 1961. Beitrage zur Kenntnis der Iranischen Binnengewasser. II. 

Regional-limnologische Studie mit besonderer Beriicksichtigung der 

Crustaceenfauna. Internationale Revue der Gesamten Hydrobiologie 46: 

309^06. 
Lucks, R. (1929). Cyclops phaleratus, Koch. Ein Beitrag zu einer 

Entwicklungsgeschichte. Bericht des Weslpreussichen Botanisch-Zoologisehen 

Vereins SI: 9-33. 
Manfredi, P. 1923. Etude sur le developpement larvaire de quelques especes du 

genre Cyclops. Annates de Biologie Lacustre 12: 272-303, 2 pis. 
Mazepova, G.F. 1962. Donnye tsilklopy yuzhnogo Baikala. In. Systematika i 

ekologiya rakoobraznykh Baikala. Trudy limnologicheskoi Instituta, Akademyia 

Nauk 2: 172-195. 
Mazepova, G.F. 1978. Cyclopoids of Lake Baikal. Trudy Limnologicheskogo 

Instituta. Akademyia Nauk 28: 1-143. 
Pesce, G.L. & Galassi, D.P. 1987. Copepodi Di Acque Sotterranee Delia Sicilia. 

Animalia 14: 193-235. 
Reid, J.W. 1987a. Some Cyclopoid and Harpacticoid Copepods from Colombia. 

including descriptions of three new species. Proceedings of the Biological Society 

of Washington 100: 262-271. 
Reid, J.W. 1987b. The Cyclopoid Copepods of a wet campo marsh in central 

Brazil. Hydrobiologia 153: 121-138. 
Strayer, D. 1988. New and rare Copepods (Cyclopoida and Harpacticoida) from 

freshwater interstitial habitats in southeastern New York. Stygologia 4: 279-29 1 . 



Bulletin of The Natural History Museum 
Zoology Series 



Earlier Zoology Bulletins are still in print. The following can be ordered from Intercept (address on inside front cover). Where the complete backlist is 
not shown, this may also be obtained from the same address. 



Volume 53 

No. 1 Puellina (Bryozoa: Cheilostomata: Cribrilinidae) from 

British and adjacent waters. J. D. D. Bishop & B. G. 

Househam. 1987. Pp. 1-63. £17.20 

No. 2 Miscellanea. 

Notes on Atlantic and other Asteroidea 5. Echinasteridae. 
Ailsa M.Clark. 

Observations on the marine nematode genus Spirina 
Gerlach, 1963 (Desmodoridae: Spiriniinae) with 
descriptions of two new species. J. W. Coles. 
Caleupodes, a new genus of eupodoid mite (Acari: 
Acariformes) showing primary opisthosomal segmentation. 
A. S. Baker. 

The Burbus perince-Barbus neglectus problem and a review 
of certain Nilotic small Barbus species (Teleostei, 
Cypriniformes, Cyprinidae). K. E. Banister. 1987. Pp. 
65-138. £20.55 

No. 3 The genera of pelmatochromine fishes (Teleostei, Cichlidae). 
A phylogenetic review. P. H. Greenwood. 1987. £17.20 

No. 4 Certain Actiniaria (Cnidaria, Anthozoa) from the Red Sea 
and tropical Indo-Pacific Ocean. K. W. England. 1987. Pp. 
205-292. £23.95 

Volume 54 

No. 1 The cranial muscles and ligaments of macrouroid fishes 
(Teleostei: Gadiformes) functional, ecological and 
phylogenetic inferences. G. J. Howes. 1988. Pp. 1-62. £16.90 

No. 2 A review of the Macrochelidae (Acari: Mesostigmata) of the 
British Isles. K. H. Hyatt & R. M. Emberson. 1988. Pp 
63-126. £17.20 

No. 3 A revision of Haplocaulus Precht, 1935 (Ciliophora: 

Peritrichida) and its morphological relatives. A. Warren. 
1988. Pp. 127-152. £8.50 

No. 4 Echinoderms of the Rockall Trough and adjacent areas. 3. 
Additional records. R. Harvey, J. D. Gage, D. S. M. Billett, 
A. M. Clark, G. L. J. Paterson. 1988. Pp. 153-198. £14.00 

No. 5 A morphological atlas of the avian uropygial gland. D. W 

Johnston. 1988. £16.60 

No. 6 Miscellanea. 

A review of the Copepod endoparasites of brittle stars 

(Ophiuroida). G A. Boxshall. 

A new genus of tantulocaridan (Crustacea: Tantulocarida) 

parasitic on a harpacticoid copepod from Tasmania. G. A. 

Boxshall. 

Unusual ascothoracid nauplii from the Red Sea. G. A. 

Boxshall & R. Bottger-Schnack. 

New nicothoid copepods (Copepoda: Siphonostomatoida) 

from an amphipod and from deep sea isopods. G. A. 

Boxshall & K. Harrison. 

A new genus of Lichomolgidae (Copepoda: 

Poecilostomatoida) associated with a phoronid in Hong 

Kong. G. A. Boxshall & A. G. Humes. 1988. £9.00 

Volume 55 

No. 1 Miscellanea. 

Structure and taxonomy of the genus Delosina Wiesner, 
1931 (Protozoa: Foraminiferida). S. A. Revets. 
Morphology and morphogenesis of Parakahliella haideri 
nov. spec. (Ciliophora, Hypotrichida). H. Berger & W 



Foissner. 

Morphology and biometry of some soil hypotrichs 

(Protozoa, Ciliophora) from Europe and Japan. H. Berger 

& W Foissner. 

Polyclad turbellarians recorded from African waters. S. 

Prudhoe, O.B.E. 

Ten new taxa of chiropteran myobiids of the genus 

Pteracarus (Acarina: Myobiidae). K.. Uchikawa. 

Anatomy and phylogeny of the cyprinid fish genus 

Onychostoma Gunther, 1896. C. Yiyu. 1989. Pp. 

1-121. £30.00 

No. 2 Studies on the Deep Sea Protobranchia: The Subfamily 
Ledellinae (Nuculanidae). J. A. Allen & F. J. Hannah. 
1989. £38.00 



Volume 56 

No. 1 Osteology of the Soay sheep. J. Clutton-Brock, K. 
Dennis-Bryan, P. L. Armitage & P. A. Jewell. 
A new marine species of Euplotes (Ciliophora, 
Hypotrichida) from Antarctica. A. Valbonesi & P. Luporini. 
Revision of the genus Eizalia Gerlach, 1957 (Nematoda: 
Xyalidae) including three new species from an oil producing 
zone in the Gulf of Mexico, with a discussion of the sibling 
species problem. D. Castillo-Fernandez & P. J. D. 
Lambshead. 

Records of Neba'.ia (Crustacea: Lepostraca) from the 
Southern Hemisphere a critical review. Erik Dahl. 
1990. £31.00 

No. 2 Tinogullmia riemanni sp. nov. ( Allogromiina: 

Foraminiferida), a new species associated with organic 

detritus in the deep-sea. A. J. Gooday. 

Larval and post-larval development of Anapagurus 

chiroacanthus (Lilljeborg, 1855) Anomura: Paguroidea: 

Paguridae. R. W Ingle. 

Redescription of Martialia hyadesi Rochebrune and 

Mabille, 1889(Mollusca: Cephalopoda) from the Southern 

Ocean. P. G. Rodhouse & J. Yeatman. 

The phylogenetic relationships of salmonoid fishes. C. P. J. 

Sanford. 

A review of the Bathygadidae (Teleostei: Gadiformes). G. J. 

Howes & O. A. Crimmen. 1 990. £36.00 



Volume 57 

No. 



No. 2 



Morphology and biometry of twelve soil testate amoebae 
(Protozoa, Rhizopoda) from Australia, Africa and Austria. 
G. Liiftenegger & W Foissner 

A revision of Cothurnia (Ciliophora: Peritrichida) and its 
morphological relatives. A. Warren & J. Paynter 
Indian Ocean echinoderms collected during the Sinbad 
Voyage (1980-81): 2. Asteroidea. L. M. Marsh & A. R. G. 
Price 

The identity and taxonomic status of Tilapia arnoldi 
Bilchrist and Thompson, 1917 (Teleostei, Cichlidae). P. H. 
Greenwood 

Anatomy, phylogeny and txonomy of the gadoid fish genus 
Macruronus Gunther, 1 873, with a revised hypothesis of 
gadoid phylogeny. G. J. Howes £38.50 

The pharyngobranchial organ of mugilid fishes; its structure, 
variability, ontogeny, possible function and taxonomic 
utility. I. J. Harrison & G. J. Howes 
Cranial anatomy and phylogeny of the South-East Asian 



catfish genus Belodontichthys. G. J. Howes & A. Fumihito 

A collection of seasnakes from Thailand with new records 

of Hydrophis belcheri (Gray). C. J. McCarthy & D. A. 

Warrell 

The copepod inhabitants of sponges and algae from Hong 

Kong. S. Malt 

The freshwater cyclopoid of Nigeria, with an illustrated key 

to all species. G. A. Boxshall & E. I. Braide 

A new species of Ferdina (Echinodermata: Asteroidea) from 

the Sultanate of Oman with discussion of the relationships 

of the genus within the family Ophidiasteridae. L. M. Marsh 

& A. C. Campbell £38.50 

Volume 58 

No. 1 The morphology and phylogeny of the Cerastinae 
(Pulmonata: Pupilloidea). P. B. Mordan 
A redescription of the uniquely polychromatic African 
cichlidfish Tilapia guinasana Trewavas, 1936. P. H. 
Greenwood 

A revision and redescription of the monotypic cichlid genus 
Pharyngochromis (Teleostei, Labroidei). P. H. Greenwood 
Description of a new species of Microgale (Insectivora: 
Tenrecidae) from eastern Madagascar. P. D. Jenkins 
Studies on the deep-sea Protobranchia (Bivalvia): the family 
Nuculidae. P. M. Rhind and J. A. Allen £40.30 

No. 2 Notes on the anatomy and classification of ophidiiform 
fishes with particular reference to the abyssal genus 
Acanthonus Gunther, 1878. G. J. Howes 
Morphology and morphogenesis of the soil ciliate Bakuella 
edaphoni nov. spec, and revision of the genus Bakuella 
Agamaliev & Alekperov, 1976 (Ciliophora, Hypotrichida). 
W. Song, N. Wilbert and H. Berger 
A new genus and species of freshwater crab from 
Cameroon, West Africa (Crustacea, Brachyura, 
Potamoidea, Potamonautidae). N. Cumberlidge and P. F. 
Clark 

On the discovery of the male of Mormonilla Giesbrecht, 
1891 (Copepoda: Mormonilloida) R. Huys, G. A. Boxshall 
and R. Bottger-Schnack £40.30 

Volume 59 

No. 1 A new snake from St Lucia, West Indies. G. Underwood 

Anatomy of the Melanonidae (Teleostei: Gadiformes), with 
comments on its phylogenetic relationships. G. J. Howes 
A review of the serranochromine cichlid fish genera 
Pharyngochromis, Sargochromis, Serranochromis and Chetia 
(Teleostei: Labroidei). P. H. Greenwood 
A revision of Danielssenia Boeck and Psammis Sars with the 
establishment of two new genera Archisenia and 
Bathypsammis (Harpacticoida: Paranannopidae). R. Huys 
and J. M. Gee. 



A new species of Syrticola Willems & Claeys, 1982 
(Copepoda: Harpacticoida) from Japan with notes on the 
type species. R. Huys and S. Ohtsuka 
Erratum £40.30 

No. 2 The status of the Persian Gulf sea snake Hydrophis 

lapemoides (Gray, 1 849) (Serpentes, Hydrophiidae). A. 

Redsted Rasmussen. 

Taxonomic revision of some Recent agglutinated 

foraminifera from the Malay Archipelago, in the Millett 

Collection, The Natural History Museum, London. P. 

Bronnimann and J. E. Whittaker. 

Foregut anatomy, feeding mechanisms, relationships and 

classification of the Conoidea (=Toxoglossa)(Gastropoda). 

J. D. Taylor, Y. I. Kantor and A.V. Sysoev. 1993. Pp. 

97-???. £40.30 

Volume 60 

No. 1 A new subfamily and genus in Achatinidae (Pulmonata: 
Sigmurethra). A. R. Mead. 

On Recent species of Spiraserpula Regenhardt, 1961 , a 
serpulid polychaete genus hitherto known only from 
Cretaceous and Tertiary fossils. T Gottfried Pillai and H.A. 
Ten Hove. 1994. Pp. 1-104. £40.30 

No. 2 Phylogenetic relationships between arietellid genera 

(Copepoda: Calanoida), with the establishment of three 
new genera. S. Ohtsuka, G. A. Boxshall and H. S. J. Roe. 
1994. Pp. 105-???. £40.30 

Volume 61 

No. 1 A revised familial classification for certain cirrhitoid genera 
(Teleostei, Percoidei Cirrhitoidea), with comments on the 
group's monophyly and taxonomic ranking. PH. 
Greenwood. 

Studies on the deep-sea Protobranchia (Bivalvia); the 
Subfamily Yoldiellinae. J.A. Allen, H.L. Sanders and F. 
Hannah. 1995. Pp. 1-90. £40.30 

No. 2 Primary studies on a mandibulohyoid 'ligament' and other 
intrabucal connective tissue linkages in cirrhitid, latrid and 
cheilodactylid fishes (Perciformes: Cirrhitoidei). PH. 
Greenwood 

A new species of Crocidura (Insectivora: Soricidae) 
recovered from owl pellets in Thailand. RD. Jenkins and 
A.L. Smith 

Redescription of Sudanonautes Floweri(De Man, 1901) 
(Brachyura: Potamoidea: Potamonautidae) from Nigeria 
and Central Africa. N. Cumberlidge 
Association of epaxial musculature with dorsal-fin 
pterygiophores in acanthomorph fishes, and its phylogenetic 
significance. R.D. Mooi and A.C. Gill. 1995. Pp. 
91-138. £40.30 



GUIDE FOR AUTHORS 



Policy. The Bulletin of the British Museum (Natural History) 
Zoology, was established specifically to accommodate 
manuscripts relevant to the Collections in the Department of 
Zoology. It provides an outlet for the publication of taxonomic 
papers which, because of their length, prove difficult to publish 
elsewhere. Preference is given to original contributions in 
English whose contents are based on the Collections, or the 
description of specimens which are being donated to enhance 
them. Acceptance of manuscripts is at the discretion of the 
Editor, on the understanding that they have not been submitted 
or published elsewhere and become the copyright of the Trustees 
of the Natural History Museum. All submissions will be 
reviewed by at least two referees. 

[Manuscripts. Initially three clear, complete copies should be 
submitted in the style and formal of the Bulletin. The text must be 
typed double-spaced throughout, including references, tables 
and legends to figures, on one side of A4 paper with 2.5 cm 
margins. All pages should be numbered consecutively, beginning 
with the title page as p. 1. SI units should be used where 
appropriate. 

Whenever possible a copy of the text, once the paper has been 
accepted, should also be provided on floppy disc (see below). 
Discs should only be sent after final acceptance, as papers 
generally need revision after refereeing. If it is impossible to 
provide an appropriate disc please ensure that the final 
typescript is clearly printed. 

Authors are requested to ensure that their manuscripts are in 
final format, because corrections at proof stage may be charged 
to the author. Additions at proof stage will not normally be 
allowed. Page proofs only will be sent. 

Word-processor discs. Please follow these instructions. 

1. Ensure that the disc you send contains only the final 
version of the paper and is identical to the typescript. 

2. Label the disc with the author's name, title of the paper 
and the word-processor programme used. Indicate whether IBM 
or Apple Mac (IBM preferred). 

3. Supply the file in the word-processor format; if there is a 
facility to save in ASCII please submit the file in ASCII as well. 

4. Specify any unusual non-keyboard characters on the 
front page of the hard copy. 

5. Do not right-justify the text. 

6. Do not set a left-hand margin. 

7. Make sure you distinguish numerals from letters, e.g. 
zero (0) from O; one ( 1 ) from 1 (el) and I. 

8. Distinguish hyphen, en rule (longer than a hyphen, used 
without a space at each end to signify 'and' or 'to', e.g. the 
Harrison-Nelson technique, 91-95%, and increasingly used 
with a space at each end parenthetically), and em rule (longer 
than an en rule, used with a space at each end parenthetically) 
by: hyphen, two hyphens and three hyphens, respectively. Be 
consistent with rule used parenthetically. 

9. Use two carriage returns to indicate beginnings of 
paragraphs. 

10. Be consistent with the presentation of each grade of 
heading (see Text below). 

Title. The title page should be arranged with the full title; 
name(s) of author(s) without academic titles; institutional 
address(es); suggested running title; address for correspondence. 

Synopsis. Each paper should have an abstract not exceeding 200 
words. This should summarise the main results and conclusions of 



the study, together with such other information to make it suitable 
for publication in abstracting journals without change. References 
must not be included in the abstract. 

Text. All papers should have an Introduction, 
Acknowledgements (where applicable) and References; Materials 
and Methods should be included unless inappropriate. Other 
major headings are left to the author's discretion and the 
requirements of the paper, subject to the Editors' approval. Three 
levels of text headings and sub-headings should be followed. All 
should be ranged left and be in upper and lower case. 
Supra-generic systematic headings only should be in capitals; 
generic and specific names are to be in italics, underlined. 
Authorities for species names should be cited only in the first 
instance. Footnotes should be avoided if at all possible. 

References. References should be listed alphabetically. 
Authorities for species names should not be included under 
References, unless clarification is relevant. The author's name, in 
bold and lower case except for the initial letter, should 
immediately be followed by the date after a single space. Where 
an author is listed more than once, the second and subsequent 
entries should be denoted by a long dash. These entries should 
be in date order. Joint authorship papers follow the entries for 
the first author and an '&' should be used instead of 'and' to 
connect joint authors. Journal titles should be entered in full. 
Examples: (i) Journals: England, K..W 1987. Certain Actinaria 
(Cnidaria, Anthozoa) from the Red Sea and tropical 
Indo-Pacific Ocean. Bulletin of the British Museum (Natural 
History j. Zoology 53: 206-292. (ii) Books: Jeon, K.W 1973. The 
Biology of Amoeba. 628 p. Academic Press, New York & 
London, (iii) Articles from books: Hartman, WD. 1981. Form 
and distribution of silica in sponges, pp. 453^93. //;: Simpson, 
T.L. & Volcani, B.E. (eds) Silicon and Siliceous Structures in 
Biological Systems. Springer- Verlag, New York. 

Tables. Each table should be typed on a separate sheet designed 
to extend across a single or double column width of a Journal page. 
It should have a brief specific title, be self-explanatory and be 
supplementary to the text. Limited space in the Journal means that 
only modest listing of primary data may be accepted. Lengthy 
material, such as non-essential locality lists, tables of 
measurements or details of mathematical derivations should be 
deposited in the Biological Data Collection of the Department of 
Library Services, The Natural History Museum, and reference 
should be made to them in the text. 

Illustrations. Figures should be designed to go across single 
(84 mm wide) or double (174 mm wide) column width of the 
Journal page, type area 235 x 174 mm. Drawings should be in 
black on white stiff card or tracing film with a line weight and 
lettering suitable for the same reduction throughout, either 50%, 
30% or 25%. After reduction the smallest lettering should be not 
less than 10 pt (3 mm). All photographs should be prepared to 
the final size of reproduction, mounted upon stiff card and 
labelled with press-on lettering. Components of figure-plates 
should be abutted. All figures should be numbered consecutively 
as a single series. Legends, brief and precise, must indicate scale 
and explain symbols and letters. 

Reprints. 25 reprints will be provided free of charge per paper. 
Orders for additional reprints can be submitted to the publisher 
on the form provided with the proofs. Later orders cannot be 
accepted. 



CONTEf 

Deep-sea conolidean gastropods collected by the John Murray Expedition, 1933-34 

A. V. Sysoev 

Reassessment of 'Calcinus' astathes Stebbing 1924 (Crustacea: Anomura: Paguridea: 

Diogenidae) 

P.A, McLaughlin 

On a new species of Ophidiaster (Echinodermata: Asteroidea) from southern China 

Y.Liao and A.M. Clark 

The life cycle of Paracyclops fimbriatus (Fischer, 1853) (Copepoda, Cyclopoida) 

S. Karaytug and G.A. Boxshall 



ZOOLOGY SERIES 

Vol. 62, No. 1, June 1996 



■ ^*9E3B8&989